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1903, vol. I. 












{Elected April 29th, 1903.) 
His Grace Tue Duke op Bedford, K,G., President. 

William T. Blanfoed, Esq., 

I.Ij.D.,F.B..S., Vice-President. 
George A. Boulexgee, Esq., 

F.R.S., Vice-President. 
William E. de Winton, Esq., 

Acting -Superi7itendent of the 

Herbert Deuce, Esq., F.L.S. 
Chaeles Drummond, Esq., 

Charles H. Gatty, Esq., LL.D. 
Feedeeick Gillett, Esq. 
F. DuCane Godbian, Esq., 

D.C.L.,F.Il.S., Fice-Presiffeii. 
Albert Gunthee, Esq., M.D., 

Ph.D., F.R.S., Vice-President. 

Oapt. The Marquis of Hamil- 
ton, M.P. 

Prof. George B. Howes, D.Sc, 
LL.D., F.R.S., Vice-President. 

Lt.-Ool. L. Howard Ieby. 

Sir Edmund G. Loder, Bt. 

E. G. B. Meade- Waldo, Esq. 

P. Chalmers Mitchell, Esq., 
M.A., D.Sc, Secretary. 

Sir Thomas Paine. 

E. Lort Phillips, Esq. 

David Sharp, Esq., M.D., 

Oldfield Thomas, Esq., F.R.S. 

Henry Woodward, Esq., LL.D., 
F.R.S. , Vice-President. 


P. Chalmers Mitchell, Esq., M.A., D.Sc, Secretary. 
W. E. DE Winton, Esq., Acting-Superintendent of the 

Frank E. Beddard, Esq., M.A., F.R.S., Prosector. 
Me. F. H. Waterhouse, Librarian. 
Mr. John Barrow, Accoicntant. 
Mr. W. H. Cole, C?tief Clerk. 

Mr. George Arthur Doubleday, Clerk of Publications. 
Me. Arthur Thomson, Assistant Superintendent of the 



January 20, 1903. 


The Secretary. Ropoit on the Additions to the Society's 

Menagerie in December 1902 1 

Mr. P. L. Sclater. On the Zebra-and-Pony Hybrid living 

in the Society's Menagerie 1 

Mr. J. S. Budgett, M.A., F.Z.S. Account of his recent 
Journey to Uganda in search of the Okapi and 
Polyjyterus 2 

1. Note on the Spiracles of Polypterus. By J. S. Budgett, 

M.A., F.Z.S 10 

2. On the Brains of N'asalis larvahts and of some otlier 

Old World Primates. By Frank E. Beddard, F.R.S. 

&c 12 

3. On the Fishes collected by Mr. G. L. Bates in Southern 

Cameroon. By G. A. Boulenger, F.R.S. , V.P.Z.S. 
(Plates I.-V.) 21 

4. On the Anatomy of the Gcphyrean PJiascolosoma teres, 

n. sp. By W. K. IIuttox, M.A., M.B., Senior Demon- 
strator of Anatomy in the University of Glasgow. 
(Plates VI.-VIII.) 29 

5. On Potamon {Potamonautes) latidactylum, a new Ficsh- 

watoi' Crali from Upper Guinea. By Dr. J. G. de Man, 

of lerseke, Holland. (Plato IX.) 41 

a 2 



6. On a new " Bird's-dung " Spider from Ceylon. By R. I. 

PococK, F.Z.S., and the Hon. IST. 0. Rothschild, B.A., 
F.KS., F.Z.S. (Plate X.) 48 

7. On the Crustaceans of the Genera Petalidium and Ser- 

gestes from the ' Challenger,' with an Account of 
Luminous Organs in Sergestes chcdlengeri, n. sp. By 
Dr. H. J. Hansen (Copenhagen). (Plates XI. & XII.) 52 

February 3, 1903. 

The Secretary. Report on the Additions to the Society's 

Menagerie in January 1903 • — 79 

1. Notes on the Hair-Slope of four Typical Mammals. By 

Walter Kidd, F.Z.S 79 

2. A Prodromus of the Snakes hitherto recorded from China, 

Japan, and the Loo Choo Islands ; with some Notes. 
By Captain F. Wall, Indian Medical Service 84 

3. Note on the Wild Sheep of the Kopet-Dagh. By R. 

Lydekker 102 

4. On new Parasitic Copepoda from Zanzibar and East 

Africa, collected by Mr. Cyril Crossland, B.A., B.Sc. 
By Staff-Surgeon P. W. Bassett-Smith, R.N., F.Z.S. ... 104 

5. On the Original Home of the Tiger. By Col. C. E. 

Stewart, C.B., C.M.G., CLE 109 

6. On the Mode of Copulation of the Indian Elephant. By 

H. Slade, Conservator of Forests, Maymyo, Burma ... Ill 

7. On the Coelenterata collected by Mr. C. Crossland in 

Zanzibar. — I. Ceratella minima, n. sp. By Sydney J. 
HiCKSON, M.A., F.R.S., F.Z.S., Beyer Professor of 
Zoology in the Owens College, Manchester. (Plate 
XIIL) 113 

8. Contributions to our Knowledge of the Plankton of the 

Faeroe Channel. — No. YIII. By G. Herbert Fowler, 
B.A., Ph.D., F.Z.S 117 

9. On the Present Condition and Habits of the Elk in 

Norway. By H. J. Elwes, F.R.S 133 


Fel)riiary 17, 1903. 

Mr, R. E. Holding. Exhibition of, and remarks upon, some 

Skulls of Mammals showing abnormal dentition 151 

1. Oa some new Species of Spiders belonging to the Families 

Flsauridce and Senoculidce ; with Cliaracters of a new 
Genus. By Frederick Pickard-Cambridgb, B.A., 
F.Z.S. (Plates Xiy. & XY.) 15i 

2. On the Marine Fauna of Zanzibar and British East Africa, 

from Collections made by Cyril Crossland in the Years 
lyul and 1902.— PolycluBta. Pai-t 1. By Cvmul Cross- 
LAXD, B.A., B.Sc. (Plates X.\L Sc XYII.) 169 

3. On the Axis, Atlas, and Proatlas in the Higher Tlierio- 

donts. By R. Broom, M.D., B.Sc, C.M.Z.S. (Plate 
XYIII.) 177 

4. A Revision of the Fishes of the Genus Triacanthus. By 

0. Tate Regan, B.A 180 

J5. On the Geographical Yariations of the Sand-Yiper, Vipera 

aumwdytes. By G. A. Boulenger, F.R.S., Y.P.Z.S. ... 185 

6. Motes on the Habits of the Hoolock. By Geo. Candler, 

M.B.Cantab 187 

March 3, 1903. 

The Secretary. Report on t]u^ Additions to tlie Society's 

Menagerie in February 1 903 191 

The Seci-etaiy. Extract from a letter from Major C. Del me 
RadcliU'e concerning skins of a Monkey and an Otter 
from Uganda 191 

Mr. F. E. Beddard, F.R.S. Exhibition of a mounted skin 

of the Greater Bird of Paradise 1 92 

Ml'. J. L. Bonhote, F.Z.S. Exhibition of, and remarks upon, 
a photogniph of Ek'pli.-ints showing considei'able gi-owth 
oi hair, and note upon the Sanskrit name of the Tiger... 192 

Prof. F. Jeffrey Bell, F.Z.S. Exhibition of, and remarks 

upon, a Holotburian of tlu' Genus Actinopi/ga 192 

1. On a new Species of Pigmy Antelope of the Genus jS'^eo- 
tra<jus from the Cameroous District, W. Africji. By 
W. E. DE WiisToN, F.Z.S. (Plate XIX.) '.. 192 


2. On the Land Operculate Mollusca collected during the 

" Skeat Expedition " to the Malay Peninsula in 1899- 
1900. By E. K. Sykes, F.Z.S. (Plate XX.) 194 

3. The Significance of the Callosities on the Limbs of the 

Equidce^ By R. Ltdekker, F.Z.S 199 

4. Note on some Remains of Struthio karatheodoris Maj. 

of the Island of Samos. By Rudolf Martin, of Basel 
University 203 

5. On a new Genus and two new Species of Earthworms 

of the Family Eudrilidce, with some Notes upon other 
African Oligochseta. By Frank E. Beddard, M.A,, 
F.R.S., F.2.S. 210 

March 17, 1903. 

Prof. Newton, F.R.S. Exhibition of, and remarks upon, 
photographs of the White Rhinoceros, taken by Mr. 
C. R. Saunders, C3I.G., in Zululand 222 

Mr. Oldfield Thomas, F.R.S. Exhibition of a skin and 
description of a new species of Monkey, Ehinopithecus 
hrelichi. (Plate XXL) 224 

Mr. Oldfield Thomas, F.R.S. Exhibition of specimens and 
description of a new species of Dixiker, Gephalophus 
ignifer 225 

1. Observations and Experiments on Japanese Long- tailed 

Fowls. By J. T. OuNNimHAM, M.A., F.Z.S. 227 

2. On some Nudibranchs from East Africa and Zanzibar. — 

Part 11. By Sir 0. Eliot, K.O.M.G., H.M. Com- 
missioner for the East Africa Protectorate, F.Z.S. 250 

3. Contributions to the Osteology of Birds. — Part VL 

Cucidiformes. By W. P. Pycraft, F.Z.S., AX.S. 
(Plate XXII.) 258 

April 21, 1903. 

The Secretary. Report on the Additions to the Society ''s 

Menagerie in March 1903 292 

L Linnoeus and Hunter on Feather-Tracts. By Henry 

Scherren, F.Z.S 292 



2. On some Mammals collected by Cajit. II. N. Dunn, 

R.A.M.C, in the Soudan. By Oldfielu Thomas, 
F.R.S., F.Z.S 294 

3. On a Collection of Turbellaria Polycladida from the Straits 

of Malacca. (Skeat Expedition, 1899-1900.) By F. F. 
Laidlaav, B.A.Cantab., Assistant Lecturer and Demon- 
strator in the Owens College. (Plate XXIII.) 301 

4. On the Phylogenetic Cause of the Transposition of the 

Testes in Mammalia : with Remai'ks on the Evolution 
of the Diaphragm and the Metanephric Kidney. By 
W. Woodland, Univei'sity College, London 319 

5. On the Geographical Distribution of Spidci's of the Oi-der 

Mygalomorplue. By R. I. Pocock, F.Z.S 340 


'age 110, ]ine 29, for Archipelago, which read Archipelago which. 



With References to the several Articles contributed by ectch. 

Bassett-Smith, Staff-Surgeon P. W., R.N., F.Z.S. 

On new Parasitic Oopepoda from Zanzibar and East 
Africa, collected by Mr. Cyril Orossland, B.A., B.Sc 104 

Beddard, Frank E., M.A., F.Pt.S., Prosector to the Society. 

On the Brains of Nasalis larvatus and of some other 
Old World Primates 12 

Exhibition of a mounted skin of the Greater Bird of 
Paradise 192 

On a new Genus and two new Species of Earthworms 
of the Family Euclrilidte, with some Notes upon other 
African Oligochreta 210 

Bell, Prof. F. Jeffrey, M.A., F.Z.S. 

Exhibition of, and remarks upon, a Ilolothurian of the 
Genus Aciinopi/ga 192 


BoNHOTE, J. Lewis, M.A., F.Z.S. 

Exhibition of, and remarks upon, a photograph of 
Elephants showing considerable growth of hair, and note 
upon the Sanskrit name of the Tiger 192 

BouLENGER, George Albert, F.R.S., V.P.Z.S, 

On the Fishes collected by Mr. G. L. Bates in Southern 
Cameroon. (Plates I.-V.) 21 

On the Geographical Variations of the Sand-Yiper, 
Vipera ammodytes 185 

Broom, Robert, M.D., B.Sc, C.M.Z.S. 

On the Axis, Atlas, and Proatlas in the Higher Therio- 
donts. (Plate XVIII.) 177 

BUDGETT, J. S., M.A., F.Z.S. 

Account of his recent Journey to Uganda in search of 
the Okapi and Polypterus 2 

Note on the Spiracles of Polypterus 10 

Cambridge, F. Pickard. See Pickard-Cambridge, F. 

Candler, George, M.B. Cantab. 

Notes on the Habits of the Hoolock 187 

Crossland, Cyril, B.A., B.Sc. 

On the Marine Fauna of Zanzibar and British East 
Africa, from Collections made by Cyril Crossland in the 
Years 1901 and 1902.— Polychseta. Part I. (Plates 
XYI. &XVII.) 169 

Cunningham, J. T., M.A., F.Z.S. 

Observations and Experiments on Japanese Long-tailed 
Fowls 227 

De Man, Dr. J. G., lerseke, Holland. 

On Potamon (Potamoncmtes) latidactylimi, a new Fresh- 
water Crab from Upper Guinea. (Plate IX. ) 41 


De Wintox, William E., Acting- Superintendent of the 
Society's Gai-dens. 

On a new Species of Pigmy Antelope of the Genus 
Reotragus from the Cameroons District, W. Africa. 
(Plate XIX.) 11)2 

Eliot, Sir Charles, K.O.M.G., H.M. Commissioner for 
the East Afiica Protectorate, F.Z.S. 
On some Nudibranchs from East Africa and Zanzibar. — 
Part II 250 

Elwes, H. J., F.R.S., F.Z.S. 

On the Present Condition and Habits of the Elk in 
Norway 133 

Fowler, G. Herbert, B.A., Ph.D., F.Z.S. 

Contributions to our Knowledge of the Plankton of the 
Faeroe Channel.— No. VIII 117 

Hansen, Dr. H. J., Copenhagen. 

On the Crustaceans of the Genera Feicdidium and 
Sergestes from the ' Challenger,' with an Account of 
Luminous Organs in Sergestes challengeri, n. sp. (Plates 
XI. &XII.) 52 

HiCKSON, Sydney J., M.A., F.R.S., F.Z.S., Beyer Professor 
of Zoology in the Owens College, Manchester. 

On the Coelenterata collected by Mr. C. Crossland in 
Zanzibar. — I. Ceratella minima, n. s\). (Plate XIII.) ... 113 

Holding, R. E. 

Exhibition of, and remarks ui^on, some Skulls of 
Mammals showing abnormal dentition 151 

HuTTON, W. K., M.A., M.B., Senior Demonstrator of 
Anatomy in the University of Glasgow. 

On the Anatomy of the Gephyrean rhascolosoma teres, 
n. sp. (Plates VI. -VIII.) 29 



KiDD, Walter, M.D., F.Z.S. 

Notes on the Hair- Slope of four Typical Mammals ... 79 

Laidlaw, F. F., B.A.Oantab., Assistant Lecturer and 
Demonstrator in the Owens College. 

On a Collection of Turbellaria Polycladida from the 
Straits of Malacca. (Skeat Expedition, 1899-1900.) 
(Plate XXIII.) 301 

Lydekker, Richard, B.A., F.R.S., F.Z.S. 

Note on the Wild Sheep of tlie Kopet-Dagh • 102 

The Significance of the Callosities on the Limbs of the 
Equidce 199 

Martin, Rudolf, of Basel University. 

Note on some Remains of Struthio karatheodoris Maj. 
of the Island of Samos 203 

Newton, Prof. Alfred, M.A., F.R.S., F.Z.S. 

Exhibition of, and remarks upon, photographs of the 
White Rhinoceros, taken by Mr. C. R. Saimders, C.M.G., 
inZululand 222 

Pick ard- Cambridge, Frederick, B.A., F.Z.S. 

On some new Species of Spiders belonging to the 
Families Pisauridm and Senoculidce ; with Characters of 
a new Genus. (Plates XIV. & XV.) 151 

PococK, R. I., F.Z.S. 

On the Geographical Distribution of Spiders of the 
Order Mygalomorphee 340 

PococK, R. I., F.Z.S., and Rothschild, Hon. N. C, B.A., 
F.E.S., F.Z.S. 

On a new "Bird's-dung" Spider from Ceylon. (Plate X.) 48 


Pycraft, William Plane, F.Z.S., A.L.S. 

Contributions to the Osteology of Birds. — Part Yl. 
Cacidlformes. (Plate XXII.) 208 

Regan, C. Tate, B.A., British Museum, Natural History. 

A Revision of the Fi.shes of the Genus Triacanthus ... 180 

RoTiiscniLD, Hon. N. C, B.A., F.E.S., F.Z.S., and Pocock, 
R. I., F.Z.S. 

On a new " Bird's-dung" Spider from Ceylon. (Plate X.) 48 

ScHERREN, Henry, F.Z.S. 

Linnreus and Hunter on Feather-Tracts 292 

Sclater, Philip Lutley, M.A., D.Sc., Ph.D., F.R.S. 

(Secretary to the Society till January 20th, 1903). 

Report on the Additions to the Society's Menagerie in 
December 1902 1 

On the Zebra-and-Pony hybrid living in the Society's 
Menagerie 1 

Sclater, William Lutley, M.A. (Secretary to the Society 
from January 21st to April 29th, 1903). 

Report on the Additions to the Society's Menagerie in 
January 1903 79 

Report on the Additions to the Society's Menagerie in 
February 1903 1^1 

Extract from a letter from IMajor 0. Delrac Radclifle 
concerning skins of a Monkey and an Otter fi'om Uganda. 191 

Report on the Additions to the Society's Menagerie in 
March 1903 292 

Slade, H., Conservator of Forests, Maymyo, Burma. 

On the Mode of Copulation of the In<lian Elephant ... HI 


Stewart, Col. 0. E., C.B., C.M.G., CLE. 

On the Original Home of the Tiger 109 

Sykes, Ernest Ruthven, F.Z.S. 

On the Land Operculate MoUusca collected during the 
' Skeat Expedition ' to the Malay Peninsula in 1899-1900. 
(Plate XX.) 194 

Thomas, Oldfield, F.R.S., F.Z.S. 

Exhibition of a skin and description of a new species 
of Monkey, Rhinopithecus hrelichi. (Plate XXL) 224 

Exhibition of specimens and description of a new species 
of Duiker, Gephalojjhics ignifer 225 

On some Mammals collected by Capt. H. N, Dunn, 
R.A.M.O., in the Soudan 294 

Wall, Captain F., Indian Medical Service. 

A Prodromus of the Snakes hitherto recorded from 
China, Japan, and the Loo Ohoo Islands ; with some Notes. 84 

Woodland, W., University College, London. 

On the Phylogenetic Cause of the Transposition of the 
Testes in Mammalia : with Remarks on the Evolution , of 
the Diaphragm and the Metanephric Kidney 319 

1903.— Vol.. I. 


Plate rage 

1. 1. Alestes intennedms. '1. Alestes opisthotauid. 3. Am-^^ 

philiua lonf/irostris 

II. 1. Lubeo anncctens. 2. Barhus tceniurus 

III. 1. Barbels jx'ogenys. 2. Barbus batesii 

IV. Microsyjiodontis batesii | 

V. 1. Anabas pleurosti(/)iia. 2. Mastacembelus schcteri . . . . ) 


MI. y Anatomy and Histology of Phascolosoma teres 29 

VIII. i 
IX. Figs 1-6. Potamon {Potamonnutes) laiidacii/luin. Figs. 

7-9. Potatnon [Potanwnaute») africanum 41 

X. PhrynaravJine rothschildi 48 

XI. 'F'l^.l. Petalidiumfoliaceum. 2. Petalidium ]ni\. ^. Scr- \ 

gestes profu7ulus. 4. S. prehensilis. 5. S. kroyeri. I ^..^ 

6. S. similis \ 

XII. Fig. 1. Sergestes arcticus. 2. jS. challerigeri J 

XIII. Ceratella minima 113 

XIV. Spiders of the Families Pisauridae and Seuoculidae .... ( -,-•, 
XV. Spidera of the Family Pitiuurida2 I 

XVI. Figs. 1, 2, 3, 4, 7, 8. Phyllochcctopterus elioti. Figs. 5, j 

6, 9. P. pictus i 109 

XVII. Phylluchcetopterus elioti J 

XVIII. Axis, Atla^i, and Proatlas of Gomplioynathua and Tri- 

rachudon 177 

XIX. Ncotragus batesi , 192 

XX. Mollusca from the Malay Peninsula 194 

XXI. lihinopithecus brelichi 224 

X.XII. Osteology of the Cuculiformcs 2o8 

XXIII. Polyclads from the Straits of Malacca 301 

1903.— Vol. L 


1. HyLrid between Equus burchelli, c?, and E. caballus, $ 2 

2. Upper surface of cerebral hemispheres of Nasalis 13 

3. Another brain of the same species 13 

4. Longitudinal median section of brain of Nasalis 15 

5. Upper surface of cerebral hemispheres of Colobtis guereza 17 

6. Longitudinal median section of brain of Colobus guereza 17 

7. Lateral view of brain of Cynopithecus 7iiger 19 

8. Longitudinal median section of brain of Cynopithecus niger .... 19 

9. Tentacular wreath of («) Phascolosoma and (b) Phyniosoma .... 35 

10. Front view of skull and horns of adult Ram of the Kopet-Dagh 

Urial 103 

11. Chondrocarpus reticulosus and Chondrocarpus sp 105 

12. Ventriculina crosslandi 107 



15. VTelsons of larvse attributed to Thysanoessa longicaudata ...... 131 


18. Horns of fully adult Elk from Trondhjem, Norway 134 

19. Horns of fully adult Elk killed in Bjorndal 134 

20. Horns of a full-grown but not adult Elk, killed in Upper 

Namdalen 135 

21. Horns of young Elk, supposed to be 3 years old 135 

22. Tracing of shed horn picked up at Mo 142 

23. Horns of a large Elk from Lithuania, in Branicki Museum .... 143 

24. Horns of a very old Elk showing degeneration 147 

25. Tracing of a cast horn found at Solem in Bangdal by Capt. 

Ferrand 148 

26. One-horned Elk in Ipswich Museum 149 

27. Side views of heads of («) Vipeirc ammodytes f. typica and {b) 

var. meridionalis 186 

28. Front view of end of snout, showing the lepidosis 186 



:2!). Slcnll of Xcntrarpia hnfcsii l'.).'> 

•'O. liigiit ieinur of Striithio havaiJieoclorig. Caudal view 204 

'31. Txv^ht iemnx oi Strutliio karatheodoris. Rostral view 204 

32. Pelvis of Struthio karatheodoris. Ventral view 20."» 

33. Pelvis of Struthio karathcodorls. Dorsal view 20(5 

34. Lateral view of the pelvis of Struthio karathcodori^ 207 

3o. Di.^section of Stuhhnnnniu michaelmmi 211 

30, Ventral surface of Bettonia lat/ariensii 214 

37. Termination of the male ellerent apparatus of Ilettoniit. liujari- 

ensis 215 

38. Spennatophore of rurcudrilus ,sp 220 

39. Eecently-killed lihinoceros siinus, adult J 223 

40. Itecently-killed Rhinoceros simus, adult c? 224 

41. Japanese Long-tailed Fowl. Cock A, photographed April 1903. 240 

42. Japanese Long-tailed Fowl. Cock Bjphotograplied April 1903. 241 

43. Left side view of stiu-uum and shoulder-girdle of CuchIhs 

cdnorus 277 

44. Left side view of sternum and shuulder-girdle of Voua rcyuaudi . 277 

45. Dorsal aspect of the pelvis of Cacomnntis merulinus 280 

40. Dorsal aspect of the pelvis of Geococcyx mcriccmus 282 

47. Side view of same 282 

48. Ventral aspect of same 282 

40. Topographical diagram showing feather-tracts and bare spaces 

in schematic bird , 294 

50. Eye-spots of Notoplana evansii 303 

51. Male orji'ans of Notoplana ei-am^ii 304 

52. Genital apparatus of Leptoplana malaijana 300 

53. " Brain-eyes " of Scmonia peinnif/cnais 30i) 

54. Genital apparatus of Bcrrieiidalia anomala 311 

55. Eye-spots of arr/us 313 

50. Eye-spots of Prosthiostovm/n pal/idmn 317 

57. Diagram illustrating Testis Descent 334 

58. Map to illustrate the Geographical Distribution of tlie Macro- 

thelean genera of Dipluridca 344 

59. Map to illustrate the Geographical Distribution of the Atypidce, 

Brachyhothriidcr, and Mecicohotliriidrf 347 

60. Map to illustrate tlie Geographical Distribution of the C'fenizidcs 

and Miyidce 351 

61. Map to illustrate the Gei>graphical Distribution of tin; Avicu- 

lariidce 355 

Proc. Zuol. Sue. — 1903, Vol. I. 




Acolischnus (Aracbn.) 362 

Asthenoccros (Vermes) 315 

Bergendalia (Vermes) 310 

Ceratophyllidia (Moll.) 250 

Chondrocarpiis (Copep.) 104 

Microsynodontis (Pisces) 26 

Notoplana (Vermes) 302 

Paradossenus (Arachn.) 153, 155 

Pleurophyllidiella (Moll.) 251 

Ventriculina (Copep.) 106 




1903, Vol. I. (January to April). 

January 20, 1903. 

Prof. G. B. Howes, D.Sc, LL.D., F.R.S., Vice-President, 
in the Chair. 

The Secretary read the following report on the additions to the 
Society's Menagerie during the month of December 1902 : — 

The I'egistered additions to the Society's Menngerie dui'ing the 
month of December 1902 were 30 in number. Of these 15 were 
acquired by presentation, 1 was born in the Gardens, and 10 were 
received on deposit and 4 in exchange. The total number of 
departui-es dui-ing the same period, Ijy death and removals, 
was 141. 

Amongst the additions worthy of notice are two very fine speci- 
mens of the One-wattled CtiHf^ov/iuy (Oasaarms uniapjiendicidatus), 
from New Guinea, deposited by the Hon. Walter Rothscliihl, 
M.P., F.Z.S., on Dec. 30th. 

Mr. P. L. Sclater read an exti'act from a letter from Messrs. 
Stagmann, Esselen & Roos, of Pretoi'ia, dated Dec. 8th, 1902, and 
addressed to Major W. H. Birkbeck of the Remount Department, 
Johaiinesberg, fi'om whicli it appeared tJiat the hylnid Zebra, 
placed inider the Society's cai-e liy tlie King on July IDtli. 1902 

Proc. Zool. Soc— 1903, Vol. I. No. I. 1 


[Jan. 20, 

(see P. Z. S. 1902, vol. ii. p. 225), was the progeny of a male 
Zebra and a pony mare. Major Birkbeck added the following 

Text-fig. 1. 

Hybrid between Eqtms hurclielU, $ , and IS. cahallus, !j? . 

particulars: — The hybrid was rising 12 months in July 1900 
when he came in. He went home with a pony of which he was 
very fond, and if he gets troublesome the company of a quiet 
pony will pacify him at once. We gelded him, thinking Her 
Majesty might use him, and a stallion hybrid is always a terror. 
The late owner Erasmus has tried to claim compensation, but as 
the beast was a fair capture he has no case at all. At the 
surrender the Boers kept any horses of ours they had and we, of 
course, keep theirs. 

Mr. J. S. Budgett, M.A., F.Z.S., gave an account of his recent 
journey to Uganda and return by the Nile, which was illustrated 
by a large series of photographs taken by him as he went along. 

Mr. Budgett made the following remarks : — 

The special object of my journey to Uganda was to continue 
investigations on the life-history of African fishes and especially 

1903.] niS JOURNEY TO UGANDA. 3 

of Polypteriis. Having made previous attempts to solve this 
problem iimler certain eonditions in a confined arcix, it was 
thought that by ol)serving it uiider varied conditions of latitude 
and altitude, new light might be brought to bear upon it. 

Fi'om Uganda it was possible either to work down the Congo 
from its source or to return northwards down the Nile. 

On my way to Uganda, and in Uganda., I gathered what infor- 
mation I could about the two routes. The southern end of the 
Nile Valley really lies at the foot of Ruwenzori, while a short 
journey westwards from this point would bring one into the he.ul- 
w:i.ters of the Congo. 

A special inducement to t;dve the Congo route was that infor- 
mati(»n might in this joui'uey be obtaineil a,l)o>it the new animal 
Okapiajnhnstnni, and other interesting forms, Ix^lieved to exist in 
the )Semliki Forest. 

From information gathered in Uganda, it was clear that it was 
useless to hope to meet with Okapi in British territory, and, 
moreover, I here learned that the Belgians had found the Okapi 
in large numbers in the Welle country. 

I found also, what had been veiy difficult to learn before 
leaving home, that the season of the rains and the breeding of 
Polypteriis were considerably earlier at the source of the Nile than 
they wei-e further northwards ; that the Semliki Valley was a, 
most inconvenient place at which to a permanent camp 
by the river-banks, owing chiefly to the scarcity of food; and 
that only one species of Polyptei'its was to be found there, while 
at least three species were to be found in the Nile fait.her to the 

The difficulty of taking delicate appa,ratus through the Congo 
Forest to the upper watei-s of the Congo was incompara.1)ly 
greater than to the upper Nile. 

And, lastly, it was to be borne in mind that the time of year at 
which one might hope to be successful in the main object was 
that at which it was well nigh impossible to do much in the way 
of collecting the higher Vertebrata which might be supposed to be 
of interest in the Semliki Valley ; for at this time the grass is so 
high that moving away from beaten tracks is almost impossible, 
while anything smaller than giraffi^ or elephant is seldom seen. 
Bearing these fa,cts in mind, I had little hesitation in (h^ciding to 
work down the Nile, sti'iking it at a point fa,rt]ier northwards 
than the iSemliki River, in order to take advantage of the lateness 
of the sea.son in that region. Accordingly, having fitted o>it my 
safari or cai-avan at Entebbe, I started for Butya,ba, on the west 
coast of Lake Albert, on July 10th. I had the advantage of 
starting thus along a. good roa,d which had just been cleared for 
the greater part of the way, and along which the rest-houses had 
been repaired for the convenience of the Commissioner and 
Consul-General of British East Africa, Sir Charles Eliot, who 
has recently made such a. i-emarknldy rapid journey from Entebl)e 
to Gondokoro, the frontier station of Uganda on the Upper Nile. 


4 MR. J. S. BUDGETT ON [Jan. 20, 

At Kampala I diverged, however, for a few days along the old 
road to Massindi to the eastward. 

I had with me at the start 50 men and boys and my bicycle. 
So many books have been written on Uganda, that there is 
little need to describe the scenery of these tropical highlands, 
especially as Sir Harry Johnston's wonderfully complete book is 
now in everybody's hands. 

Shortly, one may say that, on going northwards from Lake 
Victoria, forest is hardly seen after leaving Kampala. We passed 
day after day through almost endless elephant-grass, with palm- 
groves and papyrus-swamps in the lower parts. The hills are 
clothed with clumps and patches of acacia and euphorbias, 
while their summits are very frequently covered with huge 
granite boulders. There were thunderstorms and rain every 
afternoon, and for the first few days I saw little in the way of 
animal life : occasionally a Civet cat would cross the path, while 
overhead Hornbills and Plantain-eaters of various species were 
common. In the valley of the Maangia for a time we were rid - 
of the everlasting elephant-grass, and here Gohus thomasi and 
Zebra were plentiful. The Cohus thomasi of this region is some- 
what different to that met with in the valley of the Nile, the 
horns having a wider curve and being stouter and of a lighter 
colour, while the animal itself is of a larger build and has more 
brilliant markings. 

The Maangia River flows northward through wide undulating 
plains, covered at this time with hay-like grasses upwards of five 
feet in height, and dotted over with very fine acacias, of a cedar- 
of-Lebanon appearance and of richest deep-green colour. The 
grasses and bushes of the roadside teem with bird-life : as we 
rode along, the little Vidua principalis, with his dozen sombre 
wives, was a constant companion, flitting just ahead of us for 
a mile or more along the road. Likewise the Common Shrike 
of these parts [Lanius excuhitoritis) has exactly the same habit 
of driving along in flocks with a caravan as the Corvinella 
corvina of the West Coast. In the marshy parts, Scopus umbretta 
was often seen. 

Then, leaving the plain, we struck up over the Bukamva hills, 
and at the crest dived into the dense grass at the side of the road, 
to travel for four weary hours over a wretched and little-used 
track, often obliterated by the tramp of elephants, and where it was 
quite impossible to make any progress with the bicycle. Then, 
descending rapidly by swampy valleys and thick jungle, we came 
suddenly into the new road to Hoima and Butyaba. From here to 
Hoima the road crossed the steep hills which form the boundary 
between Uganda and Unyoro, passing by abundant plantations of 
bananas and through many a lovely valley, at the bottom of which 
a stream ran through the richest vegetation, the banks carpeted 
with Cannas, winter cherries, and hemlock, while overhead were 
many Pterocarptos-trees with blossoms like the Alamanda flower. 
As I did not care to get too far from my safari, I would often 


bicycle on for an hoiu-, and then sit down in one of these shady 
spots, and watch the Mouse-])irds han<,ang Hke acrobats in attitudes 
most quaint, and Sun-birds darting in and out of the great red 
blossoms of the Spathodea, while often a noisy flock of Prionops 
plumatus passed hurriedly along. In the moi-e open parts I often 
saw several pnu's of Ground Hornbills, and each time one had a blue 
throat, the other a red one. I can but think that this is a sexual 
character, though which is the female and which is the male 
I was unable to detei-mine. Other bii-ds seen were Pceocephalus 
meyeri, a species of Mct^'onyx, Irrisor erythrorhynchits, Hirundo 
7'iostica, and the handsome Snipe Rhyiichciia capensis. 

A very amusing bii-d that 1 watched was Eryihropygia ruficauda, 
which is most assiduous in its courting of the female, spreading 
its tail before her like a fan. 

In the more shady parts one might often see the butterflies, as 
I reniendier seeing them in the forests of Paraguay, covering the 
ground with large patches of coloui-, in flocks according to their 

Shortly before i"e;iching Hoima the i-iver Kaf u is ci'ossed ; 
here a netwoi-k of papyi-us-swamjis with good causeways over 
them abound in duck, geese, and kingfishers of vaiious kinds. 
Then, winding upwards, a high point is reached fi'om which the 
village of Hoima is seen, and in the veiy fai- distance one can 
make out the Blue Moimtains on the other .side of Lake Albert. 
At this high point I saw a charming little Widow- bird [Vidua 
hypocherina). Lions round this part aie plentiful and somewhat 
dangerous, as they usually ai-e in countries where game is not 
abvmdant. At Hoima 1 heard that several natives had recently 
been carried oS* by the Lions. 

From here, two days' march through the so-called Budonga 
Forest brings one to the shores of Lake Albert. This Budonga 
Forest is nothing more than rather heavily- wooded scrub. It is 
true that in the ravines and gorges there ai-e strips of real forest, 
but it is not in any way comparable with tlie I'eal forest of the 
tropics, where the sky can scarcely be seen. 

This Budonga woodland teems with herds of Elephant — I 
myself calculated that there were over 200 in one herd which 
we came across. Some of the males had enormous tusks, and 
these big fellows seem to keep slightly aloof fi'oin the rest of the 
herd. I knew that we were quite close to this hei'd, as there were 
great roadways through the jungle with cpiite fresh, smoking 
dung ; and here I first notice<l what stiuck me many times 
subsequently, that when elephant-dung falls on a pathway or 
clearing, there within half an hoiu- you will constantly find, 
heaped up all round the <lung, the earth-workings of a shrew or 
mole. What is it the shrew seeks in the dung? Is it the fly- 
larva? that have been blown upon the dung, or is it the dung 
itself ? Frequently elej)hants in these parts appear of a bright 
red colour, having covered their bodies with the dust of crushed- 
vip termite hills. 

6 MR. J. s. BUDGETT ON • [Jan. 20, 

On July 29th I looked from a high point on the road on to 
Lake Albeit, a vast sheet of glistening water, 1000 feet below, 
bordei-ed on this side with level plains of park-land, broken hei'e 
and there by lagoons and swamps, where I was to try first for the 

Of the results of my Polypterus work during this journey I 
shall say nothing hei-e ; sufiice it that I stayed down by the 
lake-side from July 30th to August 15th, trapping, netting, and 
shooting. During this time the fishes most abundantly met with 
were Hyclrocyon forskalii, Alestes haremose^ Dislichodus niloiicus, 
Laheo hosei, Bagriis hayad, Eutrojims niloticus, Synodontis nicjritus, 
Tilcqna nilotica, also very large specimens of a Citharinus. 

Lates niloticihs is fi'equently caught by the natives here 5 and 
6 feet long, usually with the spear. ProtojJterus and Polypterus 
were both obtained hei-e. The River- Toi'toise {Trionyx triunguis), 
28 inches in length, and veiy lai'ge specimens of Rana occijntalis 
were also common here. 

The common Antelopes were Cobus thomasi, Cobus defassa, 
Tragelaphios scriptus, Oiibi, and Cejjhcdophtcs ceqitaiorialis. Down 
on these lake- side flats the avifauna difters in a marked mannei- 
from that in the highlands. Hyjyhantornis cucullatus was now 
building in hundreds in the water-side bushes ; Laniai-msbarbarus 
a,nd Telep)lionus senegaltis in the low bushes, with Merops albicollis, 
LairiproGoli'iis p>ur'inireus^ Pyrotnelana flammic^is, Te.rpsiphone 
perspicillata^ and Dicrurus assimilis, wei-e the bii'ds most 
frequently met with. These birds were seldom seen in the 

Laniihs excubitorms seems to have a curious habit of giving a 
peculiar chattering call whenevei'a wounded animal is near. We 
often made of this indication when ti-acking wounded beasts, 
and I have no doubt of the truth of this fact. 

From here I sti'uck due east through the Budonga Foi-est ngain 
to the Victoria Nile. The joui-ney through this woodland counti-y 
was at this time of year most arduous, all the paths being densely 
overgrown with rank grass, while in the ravines the creepers and 
hanging lianas were a gi'oat hindi-ance to the porters. Duiing the 
four days I was in this rank jungle I saw veiy little in the way of 
animal-life except Elephants, a few Baboons, and an occasional 
Pviff- Adder, one of them 4 ft. 5 in. in length, and a few interesting 
insects — Phasmidfe and Mantidte. 

Plant-life was much moi'e interesting, and almost overwhelmiug 
with its abundance of variety and its beauty. 

The handsome Nightjar (Cosmetornis vexillarms) was often seen 
at sunset in these forest-camps. 

At length we struck the old road from Masindi to Wadelai, and 
the bicycle came into use again. At my first camp along this road 
there were large numbers of a golden-eyed black Weaver-bii-d 
(Ploceus nigerrimus), which I saw nowhei-e else. In its size and 
shape, coui'ting- and nesting-habits, it resembles very closely the 
gregarious Eyphantornis cucullatus. 


The Masiudi i-oad now made stiaiglit for a higli cdiiical hill, 
from the slioiilder of which we had this part of Afrira laiil out 
as a map l)efore us. To the south, the Budon^'a Forest; to the 
west, the north end of Lake Albert, with the valley of the White 
Nile extending northwai-ds ; and immediately Ijelow, from east to 
west, the valley of the Victoria Nile. Descending from terrace to 
terrace, we at length ari-ived at the vilLige of Fajao, just helow 
the Murchison Falls, on the 22nd of August. 

This wonderful gorge has been described by Baker, Yandeleur, 
and othei'S, and their descrijitions are no exaggeration. One looks 
down on the swirling, surging water, that, leaving the base of the 
falls, sweeps round the hill on which the old fort used to stand, 
with a feeling of utter amazement at the vast numbers of leaping 
fishes, crocodiles, and hi])popotaiiii that have found their way 
into this cul-de-sac of the Nile system. 

Here 1 continued my work with moi'e success than on Lake 
Albert, the counuouest fishes being Alestes baremose, A. macro- 
lepldotufi, Laies ullotlcas, Clarias lazara^ Tllapia zihlii. 

The natives here use enoi'uious wattle-traps, which they set in 
certain fixed spots, usually out of the main force of the current, 
and often catch very lai-ge fish in them. 

On August 29th I stai-ted again, as this is a most unhealthy 
place ; many of my men were on the sick-list, and food was getting 

Once out of the goi-ge of the Victoria Nile, we into o^ien 
rolling sa-vaunali country of grass and Borassus-palms, baobabs, 
and scrubby acacias. Then crossing seveiul rivers with difficulty, 
we ari'ived at Wadelai on September 1st. 

During this stage of the journey I noticed several birds not seen 
in this pai-t of Africa before : there was Corvlnella afjinls, Paras 
leucopterus, and several species of Capitonicke, all i-eminding me, 
as did the laiidscape, of the Gambia on the West Coast. Here 
also wei-e MeliUophaijns ballockoides, Macronyx cfoceics, Urolestes 
lequatorialis, Telephonus and Craieroptus. 

Fi'om Wadelai I sent my porters on to Nimule, about 100 miles 
distant, taking my loads and servants down the river by boat. 

After a few days' work at Wadelai, we started down the river 
on the 8th of Septendier. The sceneiy on this paii of the Nile is 
very charming, the hills in many places coming right down to the 
water's edge. Here one sees the pi'ocess of the growth of \\.\Qsud 
in every stage. Beginning with the Hoating separate plants of 
Pistia stratiotes, the seeds of, first a. small floating i-ush, then of 
the " 007)1 soof" grass, settle on and gradually bind together this 
carpet of separate plants into a floating island of grass. 8o 
a1)undant are these floating islands that often we appeared to be 
stationary, even when moving at 5 or 6 miles an houi', for all the 
visible lianks wei'e moving too. Once, however, the mass lodges 
against the stationary papyrus, it quickly becomes overgrown by 
this, and is converted into permanent sud. 

Fishing villages are numerous on this pait of the Nile. The 

8 MR. J. s. BUDGET! ON [Jan. 20, 

natives make very good traps of papyrus-grass, and also hunt the 
hippopotamus with long spears with a rope and float of ambatch- 
wood attached. Amongst other fishes caught here were Mor- 
myrops, Mormyrus, Hyperopisus, and Malapterxirus. 

Much of this way the hills reti-eat, and thei-e is nothing seen 
but grass floating and grass stationary, not even bird-life to 
i-elieve the monotony. The last 20 miles, however, before reaching 
the gai-i'ison town of Nimule it is very different. The Nile flows 
straight towaixls the mountains above Nimvile, and here widens 
into beautiful lagoons covered with water-lilies, in the foreground 
sheets of Pistia of the most vivid green, in the backgrou.nd bold 
wooded hills. Here and thei'e are rocky islands with schools of 
hippopotami basking in the sun ; Bee-eaters {Melittophagus 
jjusiU'Us), the Jacana (Parra a/ricana), and the gorgeous little 
Kingfishers [Coiythoroiis cyanostygma) abound. And then the 
Nile plunges into the great Nimule gorge, to tumble down 
cataract after cataract, breakhig up and pulverizing the floating 
vegetation, and issuing again at Fort Berkley free from sud. 

I was now getting anxious about catching the Sudan Govern- 
ment steamer, which comes up once a month to Gondokoro, and 
determined to leave the Nile and go straight overland for 
Gondokoro. The actual distance was little over 100 miles, but at 
this time of year the difliculties of travelling and crossing over 
rivers in flood were such that one could not tell at all how long 
the journey would take. After passing through the Nimule 
gorge, we came to the afliuence of the Assua with the Nile. The 
Assua was now in flood, the only way of crossing being by means 
of small rafts of ambatch-wood equal to taking one load at a time. 
None of my porters were able to swim, and all had likewise to be 
crossed on the rafts. After very neai-ly losing two men down 
the rapids, the crossing was completed after eleven hours' hard 
work. Here, again, the bird-life was diflferent. I saw many birds 
while on the march that I was unable to identify. There were 
great numbers of a Weaver-bird of brownish colour with a white 
crown, building innumerable star-like nests made of straight wiry 
grasses woven in at a tangent to the nest. There were also seen 
in these parts, for the first time, Scojitelus notatus and Crypto- 
rhina af7-a, though amongst these were not seen specimens with 
red beaks as was the case on the Gambia. 

Just below its confluence with the Assua River the Nile flows on 
two sides of a high hill ; a fact which strikes one as remarkable, 
for the two branches were mountain-torrents of very little depth 
of water. 

There we left the river, and passed through country with many 
villages and a good deal of cultivation, especially ground-nuts and 
millet. The aspect of this country of the Madis struck me as 
remarkably similar to that of the Gambia : the soil was rich and 
sandy, and the nuts produced were of great size. In some of the 
valleys we saw quantities of very fine bamboo, while many of the 
trees were almost smothered by the beautiful creeping lily 
Gloriosa superha. 

1903.] ms JOURNEY TO UGANDA. 9 

In one of these villages quite 10 per cent, of the natixes ha<l 
marked elephantiasis. They were very friendly, and provided nie 
with whate\er I wanted. After three (lays' wandering hy winding 
paths from village to village, we came back to the main path hy 
the side of the Nile, which for over sixty miles runs along close 
under a range of mountains on its western hank. From this point 
northwai'ds for some time the beautifid little Pai'rot /'aknornis 
docilis was common. The only Antelopes seen in this part of the 
journey were Cobus leii,colis and a species of Danxdiscus. The 
grass seemed to get longer and longer, and marching in the early 
morning, when the heavy dew was hanging from eveiy blade of 
grass in great di'ops, was most disjigrtx'able. 

On September 19th we reached the flouri.shing village of a well- 
known chief named Adimadi. This village was situated in a. 
hollow on the top of a high hill, with natuial i-ocky foitifications 
surrounding it, and overlooking a fertile valley to the east. On 
the heights above the village I saw considerable nmnbers of what 
appeared to be a largo red (7o/o/>?<.,s- monkey, a specimen of which 
I failed to secure. 

Long-horned cattle were plentiful here, and are probably the 
same race as the long-horned cattle of Ankoli. 

On the hill-sides were numl)ers of very fine African mahogany- 
trees (yKai(i) and spi'ingsof good water. In these trees wei-e many 
kinds of Plantain-eaters and Rollers (Coracias cavdaliis). This 
was the first place duiing the whole journey that I met with any 
Rollers. The hitherto daily rainstorms were getting less frequent, 
and the diy-season was setting in. 

Fiom here we marched thiough undulating pai'k-like country 
with small trees, to a similar isolated group of hills, with the 
village of Leju ne.stling beneath. Here, again, were fine spreading 
trees, in which were numbers of beautiful glossy Starlings {S'preo 
st(.perhus), and also the King of the Sparrows [Dmemelia dinemeUa). 
Passing down from the Leju hills again, we marched through 
country of a rather barren natiue, of i-ank grass and small 
acacias. The whole way the elephant-tracks weie very numerous, 
and we came suddenly on a herd of twelve with two old tuskers 
among them. 

The country now became more and more barren, and on 
Septend)er 22nd we reached the hills again, opposite the Belgian 
station of Redjaf. Here I saw several birds I had not seen before, 
including Jferops niihicus, Vinarjo waalia, Laniarius erythrogaster , 
and, I think, Lanius collui-io, though it may have T)een a different 

At Gondokoro I sent back all my porters and LTganda servants, 
and after a few days' work fishing itc, I started northwards, on 
the Sudan Covernment steamer ' Abuklca,' for Khai-toum, on 
Septemlier 27th. 

The first few days the steamer passes through firm banks, on 
which, notwithstanding the gi'ass, we saw several water-buck 
and some bufialo. Many small villages line the banks, while 
several old Dervish forts are passed. 

10 MR. J. S. BUDGETT ON THE [Jan. 20, 

At Canissa, about 100 miles noith of Gondokoro, I changed 
into the ' Kaibar,' the post-boat to Khartoum ; then, passing 
through the sud region in three days, we came to the mouth of 
the Sobat and the land of the Shelluks. 

On my arival at Khartoum, I set to work to get Arab fisher- 
men and sei'vants, fishing-tackle, provisions, &c., and returned in 
a few days to Fashoda. 

Here I made my final attack on the PolyjJterus problem. I had 
three species of Folyi^terus to work with, while material was 
faii'ly abundant. However, after sevei-al weeks' work, I finally 
packed up my things, and disconsolately i-eturned to England ; 
having got a good deal of side-light on the life and habits of 
PolyjJterus, having seen something of the Fauna and Flora of 
the most wonderful river in the world, but having again failed in 
my principal object — namely, to obtain the early stages in the 
development of Polyj^terus. 

In conclusion, I should like to say that, throughout the journey, 
I received at the hands of the Uganda and Sudan officials the 
most courteovis and liberal assistance on all occasions. 

The following papers were read : — 

1. Note on the Spiracles of Polypterus. 
By J. S. BuDGETT, M.A., F.Z.S. 

[Received January 19, 1903.] 

On seeing a letter in 'The Field' for November 8th, 1902, by 
Mr. Boulenger, in which he says that, after observing Polypterus 
in captivity for more than a year, he had not been able to 
learn anything concerning the use of the spiracles to Polypterus, 
I determined to go over my former observations concerning these 
structures and see whether I had by chance been mistaken as to 
their use. 

I have in captivity a pair of Polyptertis senegalus kept in an 
aquarium at a temperature of 75"^ to 80° F. They are quite 
tame and regularly take food from a fork. 

On December 2nd I watched them for one hour after feeding. 
While eating, the spiracles were repeatedly rapidly opened and 
closed, though not widely. The movement was apparently inde- 
pendent of other masticatory movements. Within the hotir each 
of the pair came to the surface three times at irregular intervals. 

1. Specimen A came to the su.rface and gulped air with the 

movith ; immediately after leaving the surface, two large 
bubbles of air were discharged from under the opercula. 
During the descent to the bottom the two spiracles slightly 
opened and from each a minute bubble of air issued. 

2. Specimen B performed the same movement, but no air was 

seen to issue from the spiracles. 


3. Specimen B repcitud the niovemciit, and small huliMes oF 

air (lid issue from tlie spiracles. 

4. Specimen A repeated tlie movement, l)iit- mi air was seen to 

i.ssue from the spiracles. 

5. Specimen B repeate<l the movement violently after some 

e.\citi'ment, and a,s it met the siu'face loklclij opened the 
spiracles, forming a triaii<fular apei'ture, one side l)ein<,' 
the »side of the liead and the two other sides being the two 
plates of bone whidi form the spiraculnr Hap; whether aii- 
pussed in or out of the spiracles was impossible to see a« 
the top of the head was out of the water. No bubbles 
passed from the spiracle during descent. 
(). Specimen A repeated the movement, the s[)iraclcs did not. 
open, and no air was seen to issue from liiem during 
On Decendjer 8th I watched them again for an hour. Puli/jituras 
A and 13 came to the surface for air 8 times, and 4 times tlie 
spiracles were wldiily opened above the surface of the water, and a 
sound produced as of the sucking in of air. 

1 have often found it convenient to kill Poli/i>l(',ran by pierinng 
the crania] roof and destroying the brain. During the opi'ration 
it is quite easy to stinuxlate the brain-centres in sucli a way that 
tlie spii'acles ai-e widely ojiened a,s described above. It is possil)le 
to stimulate continuously so that the spiracles are retained in the 
widely opened condition. I believe, then, that the spiracles are 
used to take in and to give out air from the swim-bladder. At 
certain times tlie fish rises quite slowly to tlie surface in the 
horizontal position, when it wotdd be easier for it to exdiaiige 
the air in the swim-bla.dder from the surface of the head than to 
turn its liead upwards in order to take air by the montli. By 
closing the mouth and opercula., distending the body-wall and 
ojieiiing the spiracles, 1 believe the fish is alile to inhale air, and 1 
should suppose that it expires previously dui-ing the same move- 
ment, as does Protopteras. I think it possible also that in the 
very shallow water whicli this fish frequents at certain times 
of year, it may be of use to the fisli to change the air in its 
swim-bladder in this way. I have often noticed, in changing tlie 
water in a tank in wliich numlieis of these fish are confined, that 
wlu'ii the watei- is exhausted the spiracles are frequently opened. 
The position of the spiracles almost immediately ovei' the long 
slit-like glottis is in favour of the view that they are connected 
in their functions witli the latter. They seem also to be used, as 
I at first believed, to let out the excess of air fi-om the pharynx 
after the fish has taken air into the swim-bladder, either with the 
mouth or with the spiracles. 

Observation upon these points is very difficult owing to the 
rapidity with which the movement takes place ; but the fisli I 
have been watcliing liave become very tame, after three years of 
captivity, and these movements are now more slow and much 
more easily watched. 

12 MR. F. E. BEDDARD ON THE [Jan. 20, 

2. On the Brains of Nasalis larvatns and of some other Old 
World Primates. By Fkank E. Beddard, F.R.S. &c. 

[Received January 17, 1903.] 

(Text-figures 2-8.) 

Among the types of Old World Primates not studied by Messrs. 
Kiikenthal and Ziehen ^ in their othei'wise fairly exhaustive survey 
of the cerebiul convolutions in the Apes, are Nasalis (if it be a 
distinct genus), Colobus, and Cynopitheczi^s. I take the oppor- 
tunity afforded me by the possession of brains of these thi'ee 
genera to compare their charactei's with those of other Old World 
genera. The memoir of Kiikenthal and Ziehen has aided me 
greatly in this attempt on account of the clearness and accuracy 
of their figures, many of which I have been able to verify by an 
inspection of brains of the same or allied species. I do not, 
however, always find myself able to agree with the selection of 
characters which they u:se to define the various genera of Apes 
examined by them". I cannot distinguish by any tangible 
difierences the arrangement of the fui-i'ows in the genera Macacus, 
Cercopithecus, Cercocehus, and perhaps Papio. It appears to me, 
in fact, that among the Cercopithecidse there are only two plans 
of ■ cei'ebral conformation, one confined to the Cercopithecinfe and 
the other to the Semnopithecinse. The facts which I discuss in 
the present communication are confirmatory of that view. But 
the classificatory results to which they appear to lead are not a 
little surprising, and may possibly be regarded as tending to 
throw doubt upon the use of the cerebral convolutions as an 
index of anything save physiological resemblances. 

§. The Brain of Nasalis larvatus. 

I have been able to compare the brain of the young male 
Nasalis larvatus ^ with those of two other examples, which I owe 
to the kindness of Dr. Charles Hose of Borneo, and I find that 
thei'e are, as might be expected, some slight difierences of detail 
in the arrangement of the furi'ows. The accompanying drawing 
(text-fig. 2) illustrates the superior aspect of the cerebral hemi- 
spheres of the specimen of Nasalis which forms the subject 

1 " Untersuchungen iiber die Grossliirnfurclien der Primateii," Jen. Zeitschr. f. 
Naturw. xxix. (n. s.) 1895, pp. 1-122. 

2 For instance, I find that the backward prolongation of the sulcus prsecentralis, 
whose absence is stated as a characteristic of Cercopithecus, is present in C. stairsi. 
In Ci/nocephalus mormon the parallel fissure joins the Sylvian above, so that the 
statement "a [the parallel fissure] nahert sich dem hinteren Ende von S [the 
Sylvian] um S dann im Bogen zu umkreisen " is not universal in its application. 
In the same species the furrow hitherto lettered b (= inferior occipital) is not 
" entirely confined to the lateral convexity " ; it also extends ventrally. The fissures 
u and X form a complete Y-shaped furrow as in other Old World forms. Messrs. 
Kiikenthal and Ziehen write : — " x verschmilzt gar nicht oder nur scheinbar mit u." 

3 Gratiolet (Mem. sur les Plis cerobraux, &c. pi. iv. figs. 1, 2) has figured a brain 
of this animal. 



Text-fig. 2. 


Upper suifaco otCcreliral lieiiiisplieivs of Nasalis. 

a, inferior ocx'ipital fissure; c, lateral oecipital fissure; d, intraparietal : 
e, postcentralis ; If, fissure of Rolando: S, Sylvian. 

Text-fiff. 3. 

Another brain of the same si>ccies 
Letters as in text-tis. 2. 

14 MR. F. E. BEDDARD ON THE [Jan. 20, 

of the present communication, while the internal view of one 
hemisphere is shown in another drawing (text-fig. 4, p. 15). The 
variations that I have noted in the three brains are the 
following : — 

There is some variability in the exposure on the upper surface 
of the brain of the parieto-occipital fissure. This, as will be seen 
from the drawing exhibited (text- fig. 2, p. 13), is better marked in 
a large female brain, given to me by Dr. Hose, than in tlie smaller 
male Inain (text-fig. 3, p. 13). In the thii-d brain there are only 
just traces to be observed of this fissure on the supei-ficial view of 
the brain. A furrow which I identify with the postcentralis 
superior is present here in rudiment (as it is in the Macaques and 
othei- genera) occasionally. It is best developed, but practically 
on one side only (the right), in the small male which lived in the 
Society's Gardens. In the Ini-ge female l)i'ain thei-e ai-e traces 
of the fissui-e on one side only (the right) ; the fissure is absent 
in the third brain. 

The fissure of Rolando cvirves back and joins the Sylvian 
fissure in the large female bi'ain ; it does not do so in the two 
remainiug lirains. On the left side of one brain the Sylvian and 
the parallel fissures joined supeiioi'ly, a state of aftaii-s which is 
characteristic of the brains of many monkeys. 

On the right hemisphere of one brain only the lateral occipital 
fissure was incomplete, and consisted of the lower arm only, there 
being but a faint indication of the upper arm of this Y-shaped 

These appear to me to be the principal diflferences in the sulci 
of the three brains. 

I have compai'ed them carefiilly with the brains of six species 
of SeTnnopithecus ', of which I owe two to the kindness of Dr. Hose, 
while four were extracted from the skulls of specimens which 
have lived in the Society's Gardens. 

The species are as follows : — S. maurus, S. femoralis, S. hypo- 
leucus, S. rubictmdus, S. priamus, and S. enielhis. The diflerences 
between these brains and those described by Kiikenthal and 
Ziehen ^ are slight. 

The fissure lettered ^ by the above-named authors, which is 
the presylvian of other writers, is less regular in its occurrence 
than it appears to be in JVasalis, but the difference is not sufliciently 
marked to permit of any sti-ess being laid upon the fact. In only 
one instance {S. femoralis), and on one side only (the right), did 
the Sylvian fissure join the parallel fissure above. The rarity 
of this ai-i-angement is exactly as in Nasalis. In one case only 
{S. -maurus), and also on one side only (the left), were there 
indications of the fissure of Rolando joining the Sylvian fissure. 
The rai'ity of this aii'nngement is again paralleled in Nascdis. 

1 Kiikenthal and Ziehen (Jen. Zeitschr. 1895, p. 1) refer to the literature, but they 
have not included a paper by Lankester (Quart. Journ. Sci. ii. 1865). 

- " Untersuchuno'en iiber die Grosshirnf'urchen der Primaten," Jen. Zeitschr. 1895 
p. 1. 


The relations of the paiieto-oecipital and tlie Simian fissures 
a,p[>('a.r to me to be exactly the same in S'einiioj)ithecHS as they are 
in Nasalis. As a rule, they are perfectly distinct from each 
other; but in two brains the operculum was more fully developed, 
as it is in the Macaques, and thus the two fissures appeared to 

The postcentralis I found to be always present, and on lioth 
sides, though often asymmeti'ical. 

In four Semnopithecits brains I find a sm:ill furrow running 
lietween the intraparietal and the iSylvian, which may i-epresent 
the anteiioi- fork of the parallel fissuie figured liy Kiikenthal 
and Ziehen in several Apes (and lettered a' in their figures), and 
sbited to occasionally occur in Sfimnojyithecus. The existence of 
this fissure, so far as my material allows me to state, dift'crontiatos 
the brain of Sonnopithecus from that of Nasalis. 

Dr. Elliot Smith ' has mentioned in one brain of SeimiopitliP.cus 
p.ntellasi a bifurcation of the calcai-ine fissure, such as that which 
is constant, or nearly so, in Macacus and CercopiihecHS. I have 
found this well developed on one side (the left) of a brain of 
<S'. ruhicundus, and less m;irked on the same side of a brain of 
<S'. priamiis, and on the i-ight side of a brain of S. feiaoralis. It 
may be observeil, however, that in SemiioplthecH.s the T-shaped 
calciii'ine fissui-e, when it is T-shaped, is not visible on a dorsal 
view of the undivided brain as it is in Macacus. In the largest of 
the three brains of Nasalis which I have examined, there were 
indications of the same bifurcation on the light side. 

Text-fig. 4. 

Longitudinal median section of brain of Nusnh's. 
c, calloso-marginal fissure; Ca, ealcarinc ; i.p, internal jiarieto-occipital. 

The inferior occipitid sulcus is alwnys less in the Lnngui's tli.-iii 
in the Miicaques. In one Semnnjntheciis ])ra,in (the left side of 
;S'. prianius) this furrow showed :in unusual ch.uactei-, in it 
joined the parallel fissure. The s-vme occuired cm tlie right 

• Cat. Pliysiol. Ser. R. C. S. ed. 2, vol. ii. p. 420. 

16 MR. P. E. BEDDARD ON THE [Jan. 20, 

hemisphere of a brain of S. hypoleucus. This I have not seen in 

This same brain (of S. priamus) and a brain of S. entellus 
show on both sides an unusual condition of the inferior temporal 
fissui-e, which is well developed and runs pai'allel with the Sylvian 
and parallel fissures. The more usual condition in this genus is a 
much shorter fissure which is more transverse in direction, and 
that is the condition which obtains in Nasalis. The fissure is 
not figured at all by Kiikenthal and Ziehen, but is mentioned 
as being very feebly developed. Another furrow which I find to 
vary in the Semnopithecits brains at my disposal is the calloso- 
mai'ginal sulcus ; it sometimes bends up and cuts the surface of 
the brain. In Nasalis it always does. 

§. The Brain of Colobus vellerosus. 

Broadly speaking the brain of this monkey is very like that 
of a Macaque. There are, however, a few small points of 
difference, of which one at any rate may be of some little 
significance. The resemblance is so close in most particulars that 
it is really unnecessary for me to describe the brain in detail. 
The drawing exhibited herewith (text-fig. 5, p. 17) will adequately 
prove my statement. I may, however, remark that the fissure of 
Rolando quite cuts the inter-hemispheral sulcus : that the post- 
centralis is well mai'ked on both sides : that the precen trails 
superior is recognizable and has a direction parallel with the long 
axis of the brain. On one side (the right) the Syhdan and 
parallel fissures join above, as is so common on both sides with 
the Macaques. The occipital lobe is very smooth, as is often the 
case with the Macaques, and the lateral occipital sulcus is hardly 
maiked at all. The cross-piece of the characteristically Macacine 
calcarine fissure is visible when the bi-ain is inspected from above. 
A small fissure, which I have not observed in othei- Old World 
Monkeys, is to be seen on either side, behind the Simian fissure, 
and running parallel with the longitudinal axis of the brain. 
Kiikenthal and Ziehen figure, but give no name to an apparently 
similar fissure (lettered E) in the bi'ain of Lagothrix humholdti. 
I am able to confirm their demonstration of fact. They do not 
figure the same fissure in any of the Oercopithecidse. 

I am unwilling to seem to emphasize too strongly this point of 
likeness between the African Colobus and the New Woi-ld 
Lagothrix ; but I may remind zoologists that likenesses between 
Colobus and the New World Monkeys have been pointed out. 

Moi-e important than this, however, is the light which the 
brain -structui'e of Colobus appears to throw upon its relationship 
to other Old World genera. On account of the structure of its 
stomach, and for some othei' reasons also, Colobus has been 
associated with Semno2nthecus into a subfamily Semnopithecinae, 
contrasted with the Cercopithecinte which embraces the remaining 
Ceicopithecidee. It should be plain from the statements made in 




tlie present paper, and fi-om the illustrations which support them, 
that so far <is its braiii is concerned Colohus cannot l)e placed 
in close proximity to 6'em7W2nthecus. From this point of view, 

Text-fij?. 5. 

Upper surfiice of cerebral hemisplim-os of Colohus gnereza. 
Ca, cilcariue fissure. Other letters as in text-fig. 2. 

Text-fig. 6. 

Longitudinal median section of brain of Colohus guereza. 
Letters as in text-fig. 4. 

Colohus is most emphatically to be placed among the Cercopithecin?e. 
In fact, it is probably safer to follow those zoologists who do not 
subdivide the family Cercopithecidfe at all. 

Proc. Zool. See— 1903, Vol. I. No. II. 2 

18 MK. F. E. BEDDARD ON THE [Jail. 20, 

§. The Brain of Cynopithecus nigee. 

I have been able to examine a single brain of the Celebesian Ape, 
of which drawings are exhibited herewith (text-figs. 7, 8, p. 19). 

The Simian fissure runs completely across and contributes to 
a perfect operculum, as in the Macaques. But the brain of Cyno- 
pithecus differs from the brains of the latter and agrees with that of 
Semnopithecus in the fact that the infraparietal fissure of each side 
bends sharply outwards before joining the Simian fissure ; this sug- 
gests an exposure of the parieto-occipital, and a very faint groove 
(better marked upon the left side) is probably to be looked upon 
as a representative of this. The lateral occipital sulcus is a single 
fissure, that is to say, it has not the Y-shape exhibited in so many 
monkeys. There is, however, a faint depi-ession suggesting the 
upper arm of this Y . The inf ei'ior occipital sulcus is in some respects 
rather peculiar. Dr. Elliot Smith has justly pointed out ' that in 
Semnopithecus ''the inferior occipital sulcus has dwindled to most 
insignificant proportions, and unless the stuident examines a large 
series of brains he will hardly recognize in the little arc around 
the lower end of the Simian svilcus the representative of the deep 
operculated infra- occipital sulcus in the Macaques." Cynopithecus 
niger has an even smaller semicircular representative of this 
furrow, which just arches round the lateral termination of the 
Simian fissure, and is nowhere near to reaching the posterior 
extremity of the occipital lobe. Further than this, a straight 
furrow, either connected (left side) or nearly connected (right 
side) with this, runs down the temporal lobe for some distance, 
exactly parallel with and between the parallel and collateral sulci. 
This I take to be the inferior temporal sulcus of brain anatomists. 
In the Macaques and Cercopitheci this sulcus is (so far as my 
own experience goes) quite constantly represented by a short 
furrow at the lower end of the temporal lobe, and this furrow is 
a marked character of those brains, and of Ce?'cocehus a,nd Cyoio- 
cephcdus. This more ventrally placed furrow is not to be seen in 
my specimen of Cynopithecus. The existence of an upper portion of 
the inferior temporal sulcus is not, however, absolutely distinctive 
of CynajntheciLS. It is indicated in a brain of Gynocephalus 
mormon which I have at my disposal, and in Nasalis and Semno- 
pithecus. In a brain of Cynocephalus porcarius this furrow is as 
well developed as in Cyno^nthecus, but on one side only (the 
right). But, with this exception, the furrow is nowhere so fully 
developed as in Cynopithecxis. So far, therefore, it appears to be 
characteristic of this genus. It is, however, apparently impossible 
to lay much stress upon the rediiced inferior occipital sulcus of 
Cynopithecus as a point of resemblance to Se-mnopithectts in the 
absence of a lai'ge series of brains of the former genus. Messrs. 
Kiikenthal and Ziehen figure a brain of Cynocephalus sphinx in 
which the fissure is fully as reduced as it is in Cynopithecus or 
Semnopithecus. I have already point-ed out (supra, p. 12 foot- 

1 Cat. Phys. Series Mus. Roy. Coll. Suvg. vol. ii. p. 426. 




note) that this state of affairs is not universal in CynocepJudus ; 
but it has still got to be proved that it is universal in Cyno- 

Text-fiff. 7. ' 

Lateral view of brain of Cynoplthecns nU/o: 

h, inferior temporal fissure ; co, collateral ; P, parallel. 
Other letters as in te.vt-ligs. 2, 3. 

Text-fiir. 8. 

Longitudinal median section of brain of Cj/nopithecun nigcr. 
Letters as in text-fisr. 4. 

The collateral sulcus, as compared with that of the Macaques 
and Cercopitheci, is particularly well developed. It has, so to 
speak, taken advantage of the feeble developnvent of the inferior 
occipital to thrust itself forward upon the occipital lobe, where it 
is phiinly visible when the brain is viewed from behind ; it is also 
visible where it curves upwards upon the lateral aspect. It is 
the rule among Monkeys for this fissure to be largely concealed 
by the cerebellum. Its exposure for its whole length in Cyno- 
pitkecus appears to me to be a characteristic feature of the brain 
of that monkey. 



The j9«raZZe^ sulctis extends considerably beyond the Sylvian 
lis&ni'.e, and does not — as is so commonly the case with the 
Macaques — join that fissure dorsally. An interesting fact about 
this fissui'e is that it bends forward at its dorsal extremity, instead 
of being continued on in a straight line. In this bending I see 
a point <of likeness to the Semnojntheci (including Nascdis). No 
absolute distinction between Cynopithectis and Macactis can be, 
however, drawn on account of this fuii'ow, since Kiikenthal and 
^Ziehen ^figure a brain of Macacus iiiMus in which thei'e is this 
same bending forwards, and, moreover, a bifurcation of the furrow 
supei'iorly, such as I note in my example of Cynojnthecits niger. 
Hhejlsstm-e of Rolando presents no noteworthy characters ; it does 
not iTiearly cut the inter- cerebral groove. The postcentralis is 
better developed on the right side of the brain than on the left, 
and is ttransverse in both cases. The 2^recentralis superior is, on 
the other hand, better developed upon the left side than upon the 
right. It is transverse in position. Among Monkeys this fissure 
is more commonly parallel with the long axis of the brain. 

The inwdian parieto-occipitcd sidcus, visible when the brain is 
bisected longitudinally, presents what I regard as rather an 
interesting and suggestive character. This fissui^e, in the Macaques, 
&c., has a distinctly forward inclination, often making an angle 
of quite 45° with the vertical. On the other hand, in the Seonno- 
pitheci this furrow is nearly vertical (a brain of JVasalis) or with 
a distinctly backward inclination. In Cynopithecus niger this 
same furrow is nearly vertical, but with a slightly backward 
inclination, thus resembling the Semnopithecidaj more than the 

The calcarine sulctcs of Cynopithecus is not at all like that 
furrow in the brains of Cynocephalus, Macactcs, Cercopithecus, 
and Cercocebus. In the four last-named genera it is a T-shaped 
sulcus, the cross of the T appeaidng almost, sometimes in fact 
quite, upon the upper surface of the occipital lobe. This furrow is 
A'ery characteiistic of those genera. In Cynopithecus the furrow 
is simple and oblique in direction, as it is, as a general rule, among 
the Seranopithecidee. ^ 

It may be convenient to tabulate the likenesses shown in the 
brain of Cynojnthecus to that of Semnopithecus. 

The brain I'esembles that of the Semnopitheci in : — 

(1) Tlie form of the intraparietal fissure. 

(2) The backward direction of the internal parieto- occipital. 

(3) The absence of any junction between the Sylvian and 

parallel fissures. 

(4) The simple foi'm of the calcarine fissure. 

(5) The shortness of the inferior occipital fissure. 

(6) The presence of a well-marked superior portion, and the 

absence of an inferioi- portion of the inferior temporal 

The first two charactei'S are absolutely distinctive of Cyno- 



^} ;®^ 





J. Green, del. et nth. . 2., 




















^^ \ 



Itv , 


V x> 





/ tl 




J Greeru del. etlitlL . 

Mmter3a£ros .irap 


-< -h 















pithecus and Senmopithecics ; the third and fourth very nearly so, 
the last two characters are found also in C>/nocephalas, but are 
more characteristic of Cijnopithecus and Semnopithecus. 

It will be seen that if one were permitted to base a classification 
upon cerebral characters, C if no pithecus would have to be renioveil 
from its position among the Baboons and placed nearer to the 
Langurs ; this is not too extreme an interi)retation of the braiu- 
characters considered on purely morphological grounds. We may 
possil)ly regard C!/iiopit]iec>i,s as occupying a somewhat basal 
position with regard to Cynocephalus on the one hand, and 
Semiiopllhecm on the other. For, in fact, its characters occiu- 
in both, though the Uaboou-like characters are on the whole less 

3. On tlie Fishes collected by Mr. G. L. Bates in Southern 
Cameroon. By G. A. Boulenger, F.ll.S., V.P.Z.S. 

[Ivfccivud Novemboi- 28, 1902.] 

(Plates I.^V.) 

The freshwittLT iish-fauna of Cameroon is still very imperfectly 
known. A small list pviblished by Peters in 1876 ' and another 
by Lounberg in 1895'- are the only contributions that have 
hitherto appeared on this subject. Tlie collection made by Buchholz 
and reported upon by Peters was important as yielding the first 
specimen of the curious Fantodoii biochholzl, since rediscovered in 
the Niger Delta and in the Upper Congo and Ubangi. It has now 
been ascertained that this little fish flies or darts through the air, 
and is, in fact, a freshwater flying-fish. .Dr. Pellegrin, of the Paris 
IVluseum, has kindly informed me that, according to the notes of 
M. J. de Brazza, the specimen obtained in the Coiigo by this 
explorer was caught by means of a butterfly-net whilst moving 
like a dragonfly above the surface of the water. 

j\Ii-. Ct. L. Bates, whose previous collections included some very 
remarkable Batrachians described in these Proceedings, has now 
made, at my request, a rather extensive collection of freshwater 
fishes in Cameroon, of which I here give a list, together with 
descriptions of nine new species, one of which deserves to be made 
the ty[)e of a new genus. 

The specimens were obtained mostly in the Kribi River, some 
15 miles from the sea; otiiers are from a small tributary of the 
Campo River, near Efulen, Bulu Country, 1500-2000 feet; whilst 
others again are from the Mvile River, a small straim flowing 
southwards into the Campo, at about the same altitude as the 

1 Mon. Bevl. Acad. 1876, pp. 195 & 244. 

- Oihers. Vetonsk.-Ak. Forh. Stockholm, 1895, p. 179. 

22 MR. G. A. BOULENGER ON [Jan. 20, 


1. Petrocephalus simus Sauv. 


3. Marcusenius sphecodes Sauv. 

4. Marcusenius brachyhistius Gill. 


5. Brycon^thiops microstoma Gthr. 

6. Alestes longipinnis Gthr. 

7. Alestes intermedius, sp. n. (Plate I. fig. 1.) 

Depth of body 3 times in total length, length of head 4 times. 
Head as long as deep, twice as long as broad ; snout shorter than 
diameter of eye ; latter 2| to 2f times in length of head ; adipose 
eyelid indistinct ; interorbital width ^ length of head ; width of 
mouth equal to diameter of eye ; maxillary not extending quite 
to below anterior border of eye ; 14 teeth (f) in the upper jaw, 
8 in the outer row of the lower jaw ; length of lower border 
of second suborbital less than diameter of eye, Gill-rakers 
moderately elongate, 12 or 13 on lower part of anterior arch. 
Dorsal II 8, above ventrals, originating a little nearer end of 
snout than caudal, middle branched rays much produced (1| to 1§ 
as long as head) in the males. Adipose fin small, I5 to twice as 
distant from the rayed dorsal as from the caudal. Anal III 19-21, 
the outline very convex in the males. Pectoral shorter than head, 
not reaching ventral ; latter produced into a long filament in the 
males. Caudal forked. Caudal peduncle a little longer than deep. 
Scales 33-34 || , 2 between lateral line and ventral. Silvery ; a 
large black spot on the caudal peduncle and on the median rays 
of the caudal ; latter yellow, blackish at the end ; dorsal black 
and red. 

Total length 85 millim. 

Three specimens from the Kribi River. 

Intermediate between A. longipinnis Gthr. and A. thoUoni 
Pellegr, Differs from the former in the smaller scales, from the 
latter in the smaller number of anal rays. 

8. Alestes oPiSTHOTiENiA, sp. n. (Plate I. fig. 2.) 
Depth of body 34- or 3| times in total length, length of head 4 
or 4-^ times. Head a little longer than deep, twice as long as 
broad ; snout a little shorter than diameter of eye ; latter 2|- or 
3 times in length of head ; adipose eyelid very feebly developed ; 
intei'orbital width f or nearly 5 length of head ; width of mouth 
equal to diameter of eye or a little less ; maxillary not extending 
quite to below anterior border of eye ; 16 teeth (|) in the upper 


jaw, 8 in the outer row of the lower jaw ; length of lower border 
of second suborbital equal to diameter of eye. Gill-rakers 
moderately elongate, closely set, 20 to 23 on lower part of anterior 
arch. Dorsal II 8, entirely behind base of ventrals, originating a 
little nearer caudal than end of snout, twice as deep as long. 
Adipose fin small, 2| or 2| times as distant from the rayed doi-sal 
as from the caudal. Anal III 13-15. Pectoral as long as head, 
reaching ventral or nearly so far. Caudal forked. Caudal peduncle 
as long as deep. Scales 25 .t?, 2 between lateral line and ventral. 
Silvery, brownish on the back ; a black humeral spot and a black 
lateral band, commencing under the dorsal and extending on the 
median rays of the caudal ; dorsal, caudal, and ventral fins lemon- 

Total length 130 millim. 

Four specimens ; Ki-ibi Rivei- and IMvile River. 

Veiy closely allieil to A.fachsii Blgr. and A. kingsleyce Gthr. 
Distinguished from the foi-mev by the less massive foi'm and the 
longer pectoral fin ; from the latter by the more posterior position 
of tlie dorsal fin and the lai-ger eye; from botli by the more 
numei'ous gill-rakers. 



11. Xexocharax spilurus Gthr. 


12. Labeo axnectens, sp. n. (Plate II. fig. 1.) 

Body compressed, its depth nearly equal to length of head, ih 
to 5 times in total length. Head 1^- as long as broad ; snout 
rounded, strongly projecting beyond the mouth, with numerous 
nuptial tubercles ; eye supero-lateral, in the second half of the 
head, its diameter 5 to 7 times in length of head, 2| to 3| times 
in interocular width ; width of mouth, with folded lips, h to j^ 
length of head ; rostral fiap and posterior border of lip feebly 
denticulated; inner surface of lip with numerous feeble, transverse 
plica? ; a small bai'bel hidden in the folds at the sides of the 
mouth. Dorsal III 9-10, with strongly notched border' ; the 
longest ray equals the length of the head and twice that of 
the last ; the fin equally distant from end of snout and from i-oot 
of caudal. Anal 11 5 ; longest I'ay g length of head. Pectoral 
rounded, a little shorter than head, not reaching ventral ; latter 
I'eaching or nearly reaching vent, its first ray falling under the 
ninth (sixth branched) ray of the dorsal. Caudal deeply forked, 
with pointed lobes. Caudal peduncle 1)| as long as deep. Scales 
36-39|^, 3 between lateral line and ventral, 12 round caudal 
peduncle. Olive above, whitish beneath, with a rather indistinct 
darker lateral band ; fins grevish. 

Total length 210 millim. 

24 MR. G. A. BOULENGEK ON [Jan. 20, 

Two specimens from near Efnlen, 

This new species, allied to L. parvus Blgr., is interesting as 
completely connecting Laheo with Tylognathus, and showing that 
the latter genus can no longer be distinguished. 

13. Barbus progenys, sp. n. (Plate III. fig. 1.) 

Depth of body 3| times in total leng-th, length of head 
3 1 times. Snout rounded, depressed, 3| times in length of 
head ; eye supero-lateral, its diameter equal to the inter- 
orbital width and 5 times in length of head ; mouth large, its 
width 3| times in length of head, lower jaw projecting beyond 
the upper ; lips well-developed, lower interrupted on the chin ; 
barbels two on each side, anterior |^ diameter of eye, posterior as 
long as eye, the distance between them | diameter of eye. Dorsal 
III 10, last simple ray feeble, not stronger than those following, 
^ length of head ; free edge of the fin emarginate ; its distance 
from the occiput less than its distance from the caudal. Anal II 5, 
longest ray g length of head. Pectoral | length of head, not 
reaching ventral; latter slightly posterior to origin of dorsal. 
Caudal peduncle 1§ as long as deep. Scales 33^, 2| between 
lateral line and ventral, 12 round caudal peduncle. Silveiy, 
brownish on the back, dorsal and lateral scales brown at the base ; 
dorsal and anal edged with blackish, theformer with an ill-defined 
dark band across the middle ; other fins whitish. 

Total length 180 millim. 

A single specimen from the Kribi River. 

Allied to B. hotokeri Blgr., from ISTatal. Distinguished by the 
more anterior dorsal fin with 10 branched rays, the shorter anal 
fin, the narrower interorbital region. 

14. Barbus t^niurus, sp. n. (Plate II. fig. 2.) 

Depth of body 3^ to 3^ times in total length, length of head 
S| to 4 times. Snout rounded, 3| to 4 times in length of 
head ; diameter of eye 4 times in length of head, interorbital 
width 2^ to 2| times ; mouth inferior, its width 3 to 3| times 
in length of head ; lips moderately developed, interrupted on 
the chin ; barbels two on each side, anterior 1^, posterior 1| 
to 2 diameters of eye, the distance between them equal to diameter 
of eye. Dorsal III 8, last simple ray ossified and moderately 
strong, but much thicker than the fii'st branched ray, a little 
shorter than head ; free edge of the fin emarginate ; its distance 
from the occiput much less than its distance from the caudal. 
Anal III 5, longest ray f to | length of head. Pectoral f to |- 
length of head, not reaching ventral ; lattei- a little posterior to 
origin of dorsal. Caudal peduncle 1| as long as deep. Scales 
24-27 ^r-, 2i between lateral line and ventral, 12 round caudal 
peduncle. Olive-brown above, yellow on the sides and below ; a 
series of black dots on the lateral line, and a black band on each 
side of the caudal peduncle ; fins white. 


Total length 120 millim. 

Several specimens from the Krihi River and from Efulen, 
Allied to B. camj>tacanilms Blkr. and B. potamogalis Cope. 
Differs from both in the stronger third simple dorsal ray, the 
more numerous scales in the lateral line (24-27 instead of 21-24), 
and the coloration. From the first in the longer posterior barbel, 
from the second in the longer l^rbels, the smaller eye, and the 
broader interorbital region. 

15. Barbus BxiTESii, sp. n. (Plate III. fig. 2.) 

Depth of body equal to length of head, 3| times in total length. 
Snout rounded, 3 times in length of head ; diameter of eye 5 
times in length of head, interorbital width 3 times ; mouth 
inferior, its width 4 times in length of head ; lips well-developed, 
lower continuous; barbels two on each side, subequal, nearly 
twice as long as eye, the distance between them half their length. 
Doi'sal IV 8, last simple ray strong, bony, not serrated, slightly 
curved, I length of head ; fi-ee edge of the fin emarginate ; its 
distance from the occiput less than its distance from the caudal. 
Anal III 5, longest ray | length of head. Pectoral f length of head, 
not i-eaching ventral ; latter below anterior rays of doi'sal. Cai;dal 
peduncle 1| as long as deep. Scales 30 ^f, 3 between lateral line 
and ventral, 12 round caudal peduncle. Brownish above, the 
scales darker at the base, white beneath ; fins greyish. 

Total length 235 millim. 

A single specimen fi-om the Kiibi River. 

This is the fii'st discovei'ed West African representative of the 
B. hynni group. Its nearest ally is B. tanensis Gthr., from the 
Tana Pviver, East Africa. 

16. Barbus kessleri Stdr. 

17. Barbus guirali Thomin. 

18. Barilius ubangensis Pellegr. 

19. Barilius kingsley.e Blgr. 


20. Clarias liberiensis Stdr. 

21. Chrysobagrus longipinnis Blgr. 


23. Amphilius longirostris Blgr. (Plate I. fig. 3.) 
Anoplopterits longirostris Bouleng. Ann. & Mag. N. H, (7) viii. 

1901, p. 447. 

Depth of body 9 times in total length, length of head 4 times. 
Head longer than broad ; eyes small, in the second half of the 
head, twodiameters apart ; interocular width | length of snout, 

26 MR. G. A. boulejstger o^st [Jan. 20^ 

which is rounded and projects a little beyond lower jaw ; posterior 
nostril midway between eye and end of snout ; pr^maxillary teeth 
forming a very short band, measviring about ^ width of mouth ; 
maxillary barbel | length of head, reaching root of pectoral ; mandi- 
bular barbel i length of head. Dorsal I 6, nearer end of snout than 
root of caudal, first ray | length of head. Adipose short, as long 
as dorsal. Anal I 5, midway between root of ventral and root of 
caudal. Pectoral a little longer than ventral, | length of head. 
Caudal forked. Caudal peduncle 1| as long as deep. Dark olive- 
brown above, mottled with black, white beneath ; dorsal, pectorals, 
and ventrals light, with two transverse series of blackish spots ; 
caudal whitish, with some black spots, black at the base, with a 
lai'ge black blotch on each lobe. 

Total length 77 millim. 

A single specimen from hills in the Bulu country, near Efulen. 

24. Synodontis obesus Blgr. 

MiCROSYNODONTis, gen. nov. 

Closely allied to Synodontis, differing only in the absence of a 
free orbital border and of suborbital bones, in the more elongate 
form and the rounded caudal fin, and in the curious modification 
of the transverse processes of the fourth vertebi-a. 

25. MiCROSTNODONTis BATESii, sp. n. (Plate ly.) 

Depth of body 5 to 6 times in total length, length of head 4| 
to 5 times. Body subcylindrical or feebly compressed in the 
prsecaudal region, strongly compressed behind ; vent in the middle 
of the total length. Head broader than deep, 1^ to 11 as long 
as broad, without ridges or keels ; skin on vertex, occiput, and 
nuchal shield adherent to the finely rugose bones ; a small frontal 
fontanelle ; eye du-ected upwards, in middle of head, its diameter 
6 or 7 times in length of head, twice in interorbital width. Lips 
moderately developed ; maxillary barbel simple, nearly as long as 
head ; mandibular barbels with long, slender branches ; outer 
mandibular bai'bels |- length of head, inner -|. Prasmaxillaiy 
teeth small, forming a villif orm band ; mandibular teeth much 
shorter than the eye, 20 to 30 in number. Gill-cleft restricted to 
the sides. Occipito-nuchal shield a little longer than broad, 
ending in two rounded processes. Humeral process narrow, 
sharply pointed, rugose. Skin smooth. Dorsal I 6 ; spine strong, 
straight, striated but not serrated, about f length of head. Adi- 
pose dorsal low, elongate, 2 to 3 times as long as its distance fi-om 
the rayed dorsal. Anal III-IY 8-9. Pectoral spine f to | length 
of head, strong and striated, with feebly serrate outer edge, with 
12 to 17 strong retrorse teeth on the inner side. Ventral not 
reaching anal. Caudal rounded. Dark brown or blackish, with 
5 yellowish cross-bai-s above, the first on the occiput, the second 
at the base of the dorsal ; the body lower down with round 
yellowish spots ; throat and belly greyish, spotted or marbled wit' . 


dark brown ; fins vspotted with dark brown, caudal yellowish at 
the base. 

Total length 100 millim. 

Several specimens fi-om the Mvile River. 

The vei-tebral column consists of 38 vertebrse, 13 praecaudals 
and 25 caudals. The first 7 are completely united, and the 
tiunsverse processes of the 4th, which form the spring-mechanism 
in Synodontis, have a very extraordinary form. This process bears 
a spheroidal expansion in front, whilst behind, where it presses 
against the bladder, it is slightly excavated or cup-shaped. It 
may be described as similar to its homologue in /Spiodontis, but 
with a large, bell-shaped, bony knob attached to its anterior sui'face. 
The air-bladder is large, as in Synodontis. The male genital gland 
is very peculiar, being lacerated into numei'ous digitiform lobes. 


26. Haplochilus sexfasciatus Gill. 

Epiplatys sexfasciatus Gill, Proc. Acad. Philad. 1862, p. 136 
(Gaboon R.). 

I\ccilia sexfasciata Peters, Mon. Berl. Ac. 1864, p. 396 

Haplochilus infrafasciatus, part., Giinth. Cat. Fish. vi. (1866), 
pp. 313 & 357 (Old Calabar). 

Hajilocliilus sexfasciatus Giinth. 1, c. 

Lycocyprinus sexfasciatus Peters, Mon. Berl. Ac. 1868, p. 146 

Ejnplatys infrafasciatus Cope, Proc. Amer. Philos. Soc. xi, 
1871, p. 457. 

Haplochilus infrafasciatus Steind. Notes Leyd. Mus. xvi. 

1894, p. 76 (Liberia) ; Lonnberg, O^fv. Vet.-Alc. Furh. Stockh. 

1895, p. 188 (Cameroon). 

27. Haplochilus elegans Blgr. 


28. Ophiocephalus obscurus Gthr. 


29. Ax ABAS MACULATUS Thomin. 

30. AXABAS PLEUROSTIGMA, sp. n. (Plate V, fig. 1.) 

Closely related to A. kingsleyce Gthr., but snout longer, as long 
■as the eye in the adult, at least two-thirds the interorbital 
width. Dorsal XIY-XVI 10-11 ; anal VIIl-IX 10-11. Scales 
27-29 f ; lateral line 14-17/10-12. A large round blackish spot 
on the middle of the side, above the extremity of the pectoral fin ; 
no dark spot at the base of the caudal fin. 

Total length 170 millim. 

Several specimens fi'om the Kribi River, 



31. Pelmatochromis BATESII BlgT. 
Recently described fi'om tlie Benito River. 

32. Pelmatochromis subocellatus Gthr. 

33. TiLAPIA LATA Gtlir. 


34. Mastacembelus loennbergii Blgr. 

28 dorsal spines. Length of head 3| times in its distance from 

35. Mastacembelus sclateri, sp. n, (Plate Y. £g. 2.) 

Depth of body 12 to 13 times in total length, length of head 
(without rostral appendage) 6 to 7 times ; vent equally or neai-ly 
equally distant from end of snout and base of caudal ; length of 
head 2^ to 2| times in its distance from vent, and ^ to | in its 
distance from first dorsal spine. Snout 3 times as long as eye, 
ending in a trifid dermal appendage which is longer than eye ; 
cleft of mouth extending hardly to below nostril ; a strong pi'pe- 
orbital and two strong prasopercular spines. Vertical fins united 
with rounded caudal. Dorsal XXYI-XXVII 85-90 ; spines 
very short. Anal II 80-90. Pectoral not qviite ^ length of head. 
Scales very small, 19 or 20 between origin of soft dorsal and 
lateral line. Olive-brown, whitish on the belly ; a dark band on 
each side of the head, passing through the eye, sometimes 
continued on the anterior part of the body ; a more or less 
distinct series of large, dark, light-edged ocelli along the base of 
the dorsal. 

Total length 225 millim. 

This new species, named after our retiring Secretary, Mr. P. L. 
Sclater, in recognition of many favours received from him dui-ing 
his tenure of office, is based on four specimens from the Mvile 
River. 3£. sclateri differs from M. marchii Sauv. in the moi'e 
numerous doi'sal spines, from M. cri/jjtacanthics Gthr., liberiensis 
Blgr., and loennbergi Blgr. in the larger head, and from M. con- 
gicus Blgr. in the presence of only two anal spines and the still 
smaller scales. 


Plate I. 

Fig. 1. Alestes intermedhis, p. 22, natural size. 

2. Alestes opisthot tenia, p. 22, f . 

3. Avi/pliilius longirostris, p. 25, natural size. 

3 a, ,, „ Upper view of head and pectoral iin, X li'. 

3 6, „ „ Lower view of head, X H. 

Plate II, 

Fig. 1. Laheo annecfens, p. 23, f. 

2. Barbus taniurus, p. 24f, natural size. 

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ANAT M.-/ AN D T i I S '1' ]:. GY OF 


Plate III. 

Fig-. 1. Biirhus pt'oi]enys, p. 24, ^. 
2. Darhus hatesii, p. 25, ^. 

Plate IV. 

Microsiinndontis hatexii, p. 26. 

a. The whole fish, natural size. 

b. Jlouth, X 2. 

c. Skeleton, natural si/e. 

d. Upper view of skull, shonlder-trinlle, and imelial shieM, X Ih. 

e. Side view showini; air-bladder and anterior ]iart of vertebral column, X 2. 

f. Lett transverse processes of coalescent anterior vertebnu, X 3. 

Plate V. 

Fip:. 1. Atiahas plciirnstifl))ia, p. 27, ". 

2. Mastaccmbclus sclateri, p. 28, natural size. 

4. On tlio Anatomy of the Gophyrean Phai^rolosoma terfs, 
n. .sp. By "W, K. Hutton, M.A., M.B., Senior Demon- 
strator of Anatomy in the University of Glasgow. 

[Received November 6, 1902.] 
(Plates VI.-VIII.^ & Text-figure 9.) 

Some tiirte ago, while engaged in ilfedging-ojierations in the 
Fii'th of Clyde, Dr. J. F. Gemmill obtained two specimens of the 
Gephyrean worm figured in Plate VI. fig. 1. These he lianded 
over to me, and at his suggestion I undertook the task of 
identifying them. My examination having led me to the coia- 
clusion that I was dealing with an animal hitherto undescribed, 
it seemed advisable to give some account of its anatomy ; and 
this I have essayed to do in the following pages. 

Unfoi-tunately the pi-esence of sand in the alimentary tract, 
no less than the leatheiy natm-e of the animal's skin, formed aii 
almost insuperable obstacle in the way of obtaining continuous 
series of sections : as a result, my account is in some i-espects 
incomplete. The worm was dredged from a depth of 60 fathoms, 
nine miles to the south-west of the Corsewall Liglit. The bottom 
was fine mud. 

External Characters and Integuinent. 

The worm is shaped like a very long-necked Florence oil -flask 
and measures 40 mm. in length. Since, however, in botli indi- 
viduals examined the introvert was almost completely invaginated, 
in order to estimate the true length probably 8 mm. be 
added to this figure. The body of the worm is seen (Plate VI. 
fig. 2) to consist of three portions which difier externally, and are 
found upon dissection to have definite lelations to the contained 

First, occupying the anterior third or more of the animal's 

' For explanation of tlie Plates, see p. 10. 

30 MR. w. K. HUTTON ON THE [Jan. 20, 

length, and containing within it the more important organs, comes 
fa airly stout and muscular portion 2 nun. in diameter (Plate VI. 
fig. 1, A). This gradually passes posteriorly into a much slighter, 
less muscular, and longer part, which extends for more than the 
middle third of the length and measures 1 mm. in thickness ; 
while lastly, there is a short, thick, ellipsoidal piece, measuring 
5 mm. by 2"5 mm., having very thin walls which are transparent 
enough to allow the coils of gut within to be dimly seen. This 
terminal portion is quite sharply demarcated from the rest of the 
body, and forms a sack in which the coiled portion of the intestine 
(Plate YI. fig. 2, G) is contained. 

The colour of the spirit-specimens is a pale yellow over the 
anterior three-fourths of the animal's length ; on the thick 
terminal part the tint is a dirty bluish white, the contents of the 
intestine appearing through the body- wall as brownish markings. 

The skin has a slightly iridescent sheen ; it is entirely devoid 
of papillae and is of a leathery toughness. It is studded with the 
openings of innumerable cuticular glands (Plate YII. fig. 6, C.G.). 
These glands (Plate YI. fig. 3) are spheroidal, and measure '05 mm. 
in diameter. Each is lined with a single layer of rather flattened 
cells (L), and the majority are completely filled with small yellow, 
highly refractive granules (I). The glands on the introvert are 
very much smaller, and granules are not so plentiful in them as 

Though defiiiite papillae are not found, tegumentary appendages 
are represented on the introvert by chitinous hooks. These are 
ari'anged in a series of ten transverse rows (Plate YI. fig. 4), 
The individual hooks vary in length (Plate YII. fig. 5) from "06 
to "08 mm. ; they are brown in colour, almost black at the tip, are 
but slightly curved, and the isolated hooks shown in the figure 
are grooved on their concave edge for about two-thiixls of their 
length (Plate YII. fig. 5, F). Each hook has a small multicellular 
epidermal core. The first part of the introvert, that nearest the 
" head " of the animal, is perfectly smooth, and, when invaginated, 
abuts against the oviter surface of the tentacular crown through- 
out two-thirds of its length ; for the remaining distance, however, 
the tentacles touch the hook-bearing part. Laceration of the 
tentacles during evagination and invagination is averted in some 
Sipunculids by the presence of a " collar," which intervenes 
between these delicate structures and the I'ough introvert. Here 
there is no such Jprovision ; the end is gained hv a transverse 
furrowing of the skin on the hook-bearing introvert dui'ing 
invagination ; and by this means the hooks, projecting from the 
base of the groove, barely come into contact with the tentacles. 
An indication of this condition is represented in fig. 6, H.I. 
(Plate YII.), 

General Arrangement of Organs. 

Figure 2 (Plate YI.) is a drawing of a dissection of the worm. 
It has been opened along the ventral aspect, and the two retractor 


muscles (D.R., V.R.) of the left side have been cut through. The 
anterior part of the alimentary canal has been twisted aside and 
turned so that it presents to view its dorsal surface. The intro- 
vert is almost completely invaginated, the real "head" of the 
worm being about the position of the two eye-spots (E). 

The internal aspect of the body-wall has a glancing appearance 
like mother-of-peai'l ; the longitudinal muscles form a continuous 
sheet with no di\-ision into strands. Thei-e are four i-etractor 
muscles (D.R., V.R.), two dorsal and two ventral. The latter are 
more than twice the length of the former, stretching fi'om the 
termination of the anteiior muscular pai-t of the body (between 
A & B, fig. 1) to the anterior pait of the cesophagns, where they 
fuse with the doi'sal miiscles to form a sheath for that tube. They 
arise, one on either side of the nerve-cord. The dorsal muscles 
are attached to the body-wall on either side of the hind-gut, a 
little way behind the anus. In fig. 2 the only part of the 
nervous system seen is the eye-spots, which are visible in a 
dissection as a couple of black specks shining througli the walls 
of the inti'overt. 

The alimentary canal (G.F.), with the exception of the anterior 
part of the a?sophagus, is shown in full. It is, like the worm 
itself, divisible into three parts, an anterior and a posterior 
portion, comparatively straight, and a middle jiiece (G) exceed- 
ingly coiled. 

In the drawing, for the sake of clearness, that part of the gut 
(F) contained in the thin portion of the worm is shown as though 
absolutely straight ; it is really slightly crumpled. Nowhere is 
there any intestinal spiral ; thei'e is no spindle-muscle, and the 
intestine is entirely free except for about 2 mm. behind the anus, 
where the i-ectum is attached to the body-wall by thin radially- 
disposed strands of muscular fibi^e, which in sections (Plate VII. 
fig. 6, M) appear to divide the crclom into dorsal and ventral 
portions. The nephridia are noticeable objects in a dissection 
(fig. 2, B.T.) on either side of the anus. All that is visible of 
the blood-vascular system is the tortuous dorsal vessel (H). The 
generative organs are visible only in mici'oscopic prepai'ations. 


The dermo-muscular tube has the arrangement common to the 
Phascolosomidaj. Underneath the integmuent there lies a layer 
of circularly-disposed fibres (Plate VII. fig. 6, R.M.), and beneath 
this again one of fibres having a longitudinal dii'ection (L.M.). 
As already mentioned, this lattei' layer is a continuous one, witli no 
division into strands such as are seen, for example, in the genus 
Sipunculas. The musculature is thickest and sti'ongest in the 
anterior third of the animal's body ; becoming thinner as the 
body narrows, it reaches the extreme of tenuity over the ellipsoidal 
posterior part, the thinning of tlie tube appealing to take place at 
the expense of the circular layer. 

32 MR. W. K. HUTTON ON THE [Jan. 20, 

Around tlie anus and the external nephridial apertures, both 
layers furnish special sphincter-fibres ; while along either side of 
the nerve-cord, as is afterwards noted, there runs a slender longi- 
tudinal strand of muscle-fibre (N.C., fig. 6). 

The genei'al arrangement of the four special retractor muscles 
of the introvert and of the radial rectal muscular attachments 
have already been described ; there remains only, in regard to the 
former, to speak of the special appearance presented by the 
muscular fibres in this worm. On comparing the ventral 
retractors with the general longitudinal musculature, one is 
struck by the great similarity of texture presented by both. A 
glancing, bright suggestion of firmness, I'eminding one of the 
surface of fresh tendon among the higher forms, is noticeable. 
The doi'sal retractors, on the other hand, have a dull, almost a 
translucent appearance. Microscopical examination of transverse 
sections reveals little difierence between the two sets of muscles, 
save only that in a given area fewer muscular fibres are present 
in the dorsal muscles, and the amount of intei'muscular connective 
tissue appears to be greater than in similar sections of the ventral 
set (Plate VII. fig. 7, A & B). On the whole, both from the 
relatively marked shortness of the dorsal muscles and from the 
appearances just described, I incline to suspect that they cannot 
be functionally veiy active. They contain a sufiicient amount of 
connective-tissue between the proper muscular bundles to make 
the suggestion that they are in a state of metamorphosis and 
regression at least probable {vide J. Bland Sutton's ' Ligaments,' 
&c., 2nd ed., 1897, p. 3 et seq.). 

Alimentary Caned. 

Surrounding the mouth, which lies at the bottom of the depres- 
sion formed by the union of their bases, are several simple 
leaf-shaped tentacles. So far as could be ascertained by the 
examination of a series of sections, 1 5 tentacles are present, and 
of these 11 are longer than the others. Each tentacle contains a 
branch of the blood-sinus and a slender nerve (Plate VII. fig. 8). 
This figure, which represents a typical transverse section of a 
tentacle, shows that on the mesial aspect the cells covering; its 
surface are nari'ow, long, arranged in palisade form, and are 
ciliated, while those of the opposite side form a simple columnar 
epithelium. For the gi-eater part of their extent the tentacles 
ai-e free, but round the mouth their bases ai'e fused together, and 
form a series of gutters which lead to the oral apeiture. 

The mouth itself lies on the top of a papilla, whose sides are 
grooved by the above-mentioned furrows. The cells lining the 
o-rooves are cubical and ciliated, and are continuous with the 
ciliated epithelium on the mesial aspect of the tentacles. Succeed- 
ing the mouth is a short oesophagus with veiy muscular walls ; in 
section (Plate VII. fig. 10) it appears to be star-shaped, the lumen 
of the tube being encroached upon by the inward projection of a 


series of papillfe whose epithelial covering is apparently devoid of 

For the reason mentioned at the beginning of this paper, it was 
no easy niattei- to gain a cei-tain knowledge of the histology of the 
alimentary tiact. Figure 9 A (Plate VII.) shows the character of 
the epithelium of the mid-gut geneiully : lying upon the thin mus- 
cular wall are nai'row cells of varying length, with a basal nucleus 
and granular c}i;oplasm ; the outlines between the several cells 
were in many j^laces most indistinct. Tlie epitheliiun of the rectum 
(fig. 9 B) is composed of cul)ical cells, witli a distinct margin on 
the surface next the lumen of the gut. They possess cilia,, and 
were the only cells in the alimentary ti'aet in which the pr&sence 
of a ciliated boi'der could with certainty be determined. 

Circalatonj Si/stem. 

This consists of a doisal vessel, a sinus sui'rouniling the base of 
the tentacles, and tentacular vessels. 

The dorsal vessel is single, and though convoluteil it is devoid 
of diverticula : coursing along the dorsjil wall of the fore gut, it 
surrounds the base of the tentacles with a cii'cular sinus into 
which the brain dips (Plate VIII. figs. 11, 15, IG) ; from tliis 
circular vessel tentacular sinuses arise, they form noticeable 
objects in sections of the tentacles (Plate VII. fig. 8, E.8.). 

Nervous System. 

This is of the usual type, and consists of a biuin, a ventral 
nerve-chain, and a couple of sense-organs. 

The bi-ain is a small, somewhat cordiform body, measuring in its 
long axis, which corresponds with that of the animal, three-tenths 
of a millimeti'e ; it lies on the dorsal wall of the pharynx at the 
base of the tentacles ; it is suri'ounded by a fine capsule of con- 
nective-tissue, and the anterior surface in addition is covered by 
ail epithelium (Plate VIII. fig. 13, E.), which is continuous with 
that of the outer non-ciliated aspect of the tentacles ; laterally, the 
muscular tube formed l)y the union of the reti'actoivs abuts u})on it 
(fig. 15, R.M.); below the brain is the blood-sinus (B.S., figs. 11, 
l8, 15) from which the tentacular vessels arise. 

The majority of the cerebial cells are small, with a clear cyto- 
plasm and i-elatively large nuclei ; they are massed anteriorlj', 
dorsally, and postei'iorly : the central and greater part of the 
ventral aspect of the brain are composed of fibres, l)ut behind 
the point of origin of the circumresopliageal commissures there 
ai-e to be seen in the lowei- coitical pai't sevei'al very lai'ge 
ganglion-cells (G.O., fig. 11): these giant cells measure "03 mm. 
in diameter; some are pyriform, others iri'egular in shape, and 
these appear to be multipolai-, unlike those of Fhyraosoma 
(Shi])ley, No. 7.^), but on this point I cannot 1)6 absolutely 

' This iiumlicr luis vcfijn'iicc to flio list i>f iiutliovs quoted on p. '10. 

Piioc. ZooL. Sue. -1903, Vol. I. Ko. III. 3 

34 MR. W. K. HUTTON ON THE [Jail. 20, 

Fi'om the brain two pairs of nerves arise. The fii'st pair 
(Plate VIII. fig. 16, B), which are vei-y slender, take their origin 
from the ventral asj)ect of the ganglionic mass jvist anterior to the 
eye-spots ; from them twigs pass to supply the epineural canal and 
apparently also the two dorsal tentacles. The remaining tentacles 
receive their nerves from the two stout cii-cumoesophageal con- 
nectives (fig. 16, A), which constitute the second pair of nerves, 
and arise ventro-laterally behind the eye-spots, coui'sing round 
the base of the tentacular crown, between it and the muscular 
tube formed by the fusion of the retractor muscles. Yentrally, 
the cii'cumoesophageal commissures unite to form the nerve-cord. 
This shows no trace of segmentation. Dorsally it is composed of 
fibres, while ventrally nerve- cells are distributed evenly along its 
whole length. It is accompanied laterally by two fine muscles, as 
is usual in the Sipunculidte. 


In tracing back series of sections, there becomes apparent on 
the dorsal wall of the pharynx a patch of long-celled columnar pig- 
mented epithelium. The cells of this layer are found (Plate VIII. 
figs. 12, 13, 14, 15, S.E.) to be the dorsal wall of a slit- like canal, 
whose ventral wall is formed by cells which, on the one hand, cover 
the brain in fi-ont, and on the other ai'e continuous with the non- 
ciliated epithelium which clothes the outer aspect of the two 
dorsal tentacles (S.V.), which are seen in fig. 12 to have united 
along their mesial margins. 

Posteriorly the canal thus formed (which may be called the 
epineural canal) becomes very narrow centrally, and ends blindly 
by dividing into two little culs-de-sac (S.P., fig. 15), Avhich 
project downwards and outwards into the cerebiul cortex, and are 
lined by a columnar epithelium the cells of which are deeply laden 
with granules of a reddish- black pigment. 

The cup-like organs thus formed are usually called " eye-spots," 
and are present in at least four other species of Phascolosoma, but, 
so far as I am aware, no account of their structure has hitherto 
been published. Their existence in an animal which lives at a 
depth so great as that of 60 fathoms is noteworthy, and appears 
to me to ai'gue for the organs in question some function other than 
even merely photometric ; but a small fraction of difi'used daylight 
can permeate so far. They are always to be found in the genus 
Phymosoma^ and from the above-given description it will be seen 
that the sense-organs of Ph. teres resemble in the main those of 
Phymosoma varians (Shipley, 7). They differ, however, from 
those of Phymosom-a in their relations to the tentacles. 

In the genus Phymosoma the cells entering into the formation 
of the eye-spots are continuous, on the one hand, with the epithelium 
covering the anterior aspect of the brain, and, on the other, with 
those which constitute the lining of the mesial surface of the 
tentacles. In Phascolosoma, on the contrary, the sensory epi- 


thelium is furnished by a tentacular sui-face whicli is apparently 

This seeming discrepancy is readily explained by a reference to 
the following diagrams (text-fig. 9), modified from Selenka, in 
which A represents the position of the mouth, B that of the 
cerebral ganglion, while thick-dotted lines indicate the relations 
of the tentacular wreath. 

In text-fig. 9 a this wi-eath is seen surrounding the mouth with 
a simple circlet, as in Phascolosoma teres. In such a case the 
infundibuliform approach to the cerebral organ must be lined by 
cells, continuous not only with cells of the introvert epiilermis 
but with those covering the outer aspect of the tentacular 


Text-fig. 9. 


\ -s 

' \ / I B I \ 

/ \ 

/ \ A 

I A 



\ / 

a ^ 

Tentacular wri-ath of {a) FMxcolosoma and (i) FJii/mosoma. 

In text-fig. 9 b the crown of tentacles is seen to have been in- 
dented in its upper (neural) part, and the edges of the indentation 
have been bent up in such a manner as to form a hoi-seshoc- 
shaped lophophore which, partially encircling the brain, overhangs 
the mouth instead of surrounding it. The finely dotted line in the 
diagram represents the position of the "lower lip" in Phi/mosoma ; 
this may be looked upon as corresponding to the fused bases of 
aborted tentacles. Here, then, the brain, lying at the base of the 
lophophore, can only be approached by a canal whose walls are 
coated by an epithelium continuous with that covering the mesial 
suifacc of the tentacles. 

The arrangement seen in Phctscolosoma is the more primitive. 
Briefly stated, tentacular surfaces which are inner and outer in 
the genus Phascolosoma, become outer and inner respectively in 
Phymosoma, owing to the secondary indentation of the crown of 

According to Shipley {loc. cit.), the space included within the 
concavity of the lophophore of Phymosoma becomes the repre- 
sentative of the " praM)ral lol)e " of Phoronis. 

:^36 MR. W. K. nUTTON ON THE [Jan. 20, 

Altliougli nowadays, in virtue of our knowledge of the totally 
distinct modes of development of the tentacles in Phoronis and 
the Sipunculids, it does not seem possible to institute any 
homology between the lophophoral crowns of Phoronis and 
Phymosoma, yet it may be of interest to note that in the species 
of Sipunculid under consideration the homologue of the pigmented 
epithelium, which in Phymosoma covers Shipley's " prfeoral lobe," 
is to be found in that layer of pigmented cells (Plate VIII. 
fig. 12, S.E.) which forms the dorsal wall of the epineui-al canal 
as well as in that layer continuous with it, bounding the anterior 
svirface of the brain (fig. 13, E.). 


These are small (Plate A^I. fig. 2, B.T.), measuring less than the 
twentieth part of the length of the woim ; the right nephridium 
is the longer and more convoluted of the two. Each organ is 
retort-shaped (Plate YIII. fig. 17), having a bulbous portion (0) 
attached to the body-wall, and a tubular part (D) which, arising 
from the postei'ior part of the bulb, bends i-ound it mesially and 
pi'ojects freely forwards into the coelom. The wall of the bulb is 
thin, covered externally by the cells of the peritoneum (A, fig. 18), 
and intei-nally by a single layer of granular-looking columnar 
epithelium (0, fig. 18); it is composed of slender intercrossing 
muscular bands. The cavity of the bulb is packed with fine 
vesicles (E, fig. 18), which, it will be seen, are derived from the 
secreting epithelium of the tubular portion of the nephridium. 
The bvilb communicates with the exteiior by a fine canal (A, fig. 17), 
lined with a small-celled epithelium, which pierces the body- wall 
immediately in front of the anus, and its cavity is, further, 
placed in connection with the coelomic space through a slit-like 
opening (B, fig. 17) in front of the excretory canal. The lips of 
this opening, lined with ciliated epithelium, lie between the body- 
wall and the bulbous part of the nephridium, and are concealed 
by its overhanging anterior extremity. The tul^ular portion of 
the organ has a wall which is formed by a peritoneal lamella, 
beneath which is a layer of muscles (B, fig. 18). This muscular 
layer consists of a meshwork of strands, between interstices of which 
the lining-epithelium bulges outwards, forming little crypts lined 
with a single layei- of secretory cells (0). In cross section there 
are seen projecting into the lumen of the tube feathery columns 
of cells (D, fig. 18), which have an appearance comparable to that 
of goblet-cells. The nucleus is small and irregular in shape, 
placed basally ; while the cell-contents may be seen to become 
clearer near the free margin, and a.i-e evidently extruded ultimately 
as the thin-walled vesicles which are found filling the lumen of 
the organ. 

Shipley aptly compares the process, which takes place also, in 
Phymosoma, to the formation of the granules exci'eted by the 
cells of the mammaiy gland. 


The Generative Oryaus. 

Sti-etcliing across the ventral aspect of the cfeloin, at the point 
where tlie ventral retractors are attached to the hody-wall, is a 
Land of cells with relatively large nuclei. These cells const itule 
the genital hand and ai-e derived from the peritoneal epitheliuiu. 
In the single worm examined, neither free o\'a nor spermatozoa 
were to be seen in the body-cavity. 


The presence of hooks and the possession of four i-etractor 
muscles are characters which at once place rJtascolosoina teres in 
the first of the five " groups" into wliich Selenka, (5) divides the 
genus. At the ti)ue of publication of his ' Die Sipuuculiden ' three 
species alone [Ph. virjgare, Ph. elongatum, and Ph. cjjlindratmn) 
composed the gi'oup. To these must now be added, as well as 
J'h. teres, Ph. saadcri Collin (1) and Ph. lobostomum Grube 
(Fi'-clier, 2). Tlie position of Pltascolosoma ahnoriais Sluiter (ti) 
must in tlie existing state of classification remain doubtful, as the 
whole antei'ior end of the animal was wanting. 

The neai-est ally of Ph. teres in the group is Ph. elongatum 
(see Kefei-stein, 3). Both have the same arrangement of tentacles 
and the same number of rows of hooks, while the hooks themselves 
are very similar. Ph. teres, however, difl^'ers widely in the pro- 
portionate length of its introvert, and, as regards its interuiil 
anatomy, in the small size and unequal bulk of the nepluidia,, in 
the absence of any definite spiral coiling of the intestine, in the 
great length of the hind-gut, and in the absence of a spindle- 
muscle ; and, as well as in these particulars, in its characteristic 
habit of body. Fiom eveiy other member of the group except 
7Vi. ci/lindrat/i')ii it (lifters in the complete absence of papilhe. 
These peculiarities warrant the systematic zoologist in assigning 
to the animal under discussion a distinct pLice in the genus, a.nd 
I propose for it the name of Phascolosoma teres. 

That the genus Phascolosoma is an eminently variable one is 
evident to anj^ investigntor who makes a systematic study of tlie 
Sipunculidai ; and that this tendency to variation may result in 
the inclusion in one small sub-group of animals which difter in an 
ast(mishing extent in external habit, is abundantly exemplilied 
by the fact that Phascolosoma teres and Ph. sanderi (Collin) — two 
Avoi-ms so dissimilar that a. casual observer would be pai'doned for 
placing them in difterent genera — must yet, in the present rather 
luisaiisfactory state of classification, be united in the group cnlled 
by Selenka ''(iroup 1." of the Phascolosomidic ; yt^t, while tln^ 
latter was considered by its describer Collin to pi-esent points of 
i-escud)lance to that curious 8ipunculid Goljingia niacintoshii (8), 
;Mid seems to have points in common with A.y)idosiphon, the 
foniiiT appears to me to }»resent in its structui'al pecidiarities 
definite alEnities to the sedentary Phasculioii. lloule (4), who 

38 MR. W. K. HUTTON ON THE [Jail. 20, 

investigated the Phascolosomiclse obtained by the ' Ti^availleur ' 
and ' Talisman,' in speaking of classification, goes the length of 
saying, " L'espece bien affirmee n'existe pas dansle genre Phasco- 
losome " ; and by this somewhat strong assertion he would 
indicate that the genus is in such a plastic condition— in a state 
of fluctuating variability as one may term it — that it is capable 
of being influenced by what Semper called " the conditions of 
existence " in such a way, that within the genus we may have 
numbers of types, differing on the whole but little from one 
another, the links of their genetic affinity easily supplied, which 
yet point the way toward the probable origin of the other 
Sipunculid genera. As Roule expressed it : " Oe genre est encore 
en voie de transformation, il se morcelle en un grand nombre de 
types pevi dissemblables, et ne se divise point en especes precises, 
dont les variations s'enserrent dans des limites peu eloignees." 
Roule's investigations led him to the conclusion that the Phasco- 
losomidas examined by him formed a series of links connecting 
the parent Phascolosoma stock with Aspidosiphon. In the phylo- 
genetic table drawn up pi'ovisionally by Selenka in his beautiful 
monograph ' Die Sipunculiden ' (page 6), a polyphyletic origin is 
suggested for both Phascolion and Aspidosipho?i. The former 
may have arisen dii'ectly from the parent Phascolosomid, or from 
a common stem (similarly derived) which branches into Onchne- 
soma, Tylosoma, and Phascolion ; while Asjndosiphon may have 
been derived from the directly produced Phascolion, or from a stem 
common to itself, Phyniosoma, and Cloeosijyhon. 

In Phascolosoma teres we have, as it appears to me, not perhaps 
an actually intermediate type, but at the least a species which 
gives us hints as to the probable path taken in its phylogenetic 
course by Phascolion ; while the species examined by Roule play 
the same role as. regards the genus Aspidosiphon. Phascolosoma 
sanderi, though considered by Collin to approach Golfingia, has a 
distinct but superficial I'esemblance to Aspidosiphon, and was 
rightly, as I think, placed by its describer in the genus Phascolo- 

In the worm which forms the subject of this paper the follow- 
ing characters suggest Phascolion-like affinities : — The differing 
bulk of the two nephridia, the relatively feeble development of 
the dorsal retractors (a character upon which Roule placed much 
reliance in his investigations), the lack of spindle-muscle, and the 
coiled, but not spirally-wound gut, — all these are peculiarities 
which place the worm in closer relationship with Phascolion than 
• with any other of the groups derived from the original Phasco- 
losomid stock. 

Information as to the habits of Phascolosoma teres is wanting ; 
but in its mode of life it is probably sluggish, lying buried in soft 
ooze with its anterior end alone projecting. The thin- walled 
bag-like posterior portion, endowed with but feeble muscularity, 
so sharply demarcated from the rest of the body, so relatively 
large, would undoubtedly act as a hindrance to active locomotion 


througli sand oi^ mud. Most species of Phascolton live with tlio 
posterior part of tlie body either einlieiUled in mud or protected 
by some Ci\st-off mollusk-sl\ell ; and if we assume, as the external 
habit of Phascolosomn teres seems to me to warrant, this similarity 
of mode of life between the two typos, certain peculiarities of 
structure in Ph. teres become of easy explanation. 

Selenka, long ago, pointed out that those Sipuncnlids which 
live in shells usually have only one nephridium. In other wt)rds, 
in animals of sluggish habit, the needs of metabolism are served 
by one excretory tube. 

Now in Ph. teres, while there are two nophridin, they nre both 
small and ai'e of difi'ei-ent sizes ; the left has I'eti'ograded to a 
greater extent than the right. It is to be noted thsit when only 
one nephi'idium persists — as in PhascoUoa — it is that upon the 
I'ight side. 

Enfeeblement of the muscular system is correlated with inactive 
habits ; and in worms lying half buried in mini a. strongly 
developed system of retractors censes to be a clamant need. The 
extreme case is reached in Phascolion, where often the ventral 
I'eti'actoi's alone remain, while an indication of regression of the 
doi'sal i/eti'actor system is seen {vide supra) in Phascolosoma teres. 

Lastly, in this worm, the coils of intestine, completely filling 
np the sharply-delimited posterior part of the body, could scarcely 
be efficiently acted upon by any spindle-muscle, which, we see, is 

In these structund chnracters, then, we seem to oljtain in 
Ph. teres an indication of the path taken by Phascolion in its 
derivation from the more primitive Phascolosoma. 

Summary. — With the characters of the Group, 

Phascolosoma teres, n. sp. 

Body 40 mm. long, introvert ca. 8 mm. Posterior portion 
stout, sharply demarcated, ellipsoidal, containing the coiled 
portion of the gut. 

Colour yellowLsh, merging to dirty bluisli white over the 
postei'ior pait. 

Skin smooth, iiidescent, devoid of papUlfe, translucent posteriorly, 
beset with numberless glands. A simple wreath of 15 tentacles 
surrounds the mouth. Forward on the introvert 10 rows of 
backwardly directed, slightly curved, brownish hooks, •()6--08 nnu. 

Two eye-spots. Alimentary canal free, without spindle-muscle 
or mesenterial attachment save close to the anus. 8[)iral winding 
absent, but the tulie much coiled in its middle portion. Hind- 
gut very long. Two nephridia, the right the larger, one-twentieth 
of the body-length. Simple contractile vessel. 

Hab. Firth of Clyde. Dredged from 60 fiithoms. 

40 on the anatomy of a new gephyrean worm. [jan. 20, 

Works referred to. 

(1) A. Collin. — " Gephyreen gesammelt von Herrn Stabsarzt 

Dr. Sander," in Archiv fiir Naturgeschichte, Iviii. p. 197, 

(2) Fischer. — " Die Gephyreen des Naturhistorisclien Museums 

zu Hamburg," in Abhandlungen aus deni Gebiete der 
Naturwissenschaften &c., Hamburg, Bd. xiii. p. 1, 1895. 

(3) Keferstein.— " Beitrage zur Kenntniss der Gattung Phascolo- 

soma F. S. Leuck.," in Zeitschrift fiir wiss. Zoologie, 
Bd. xii. p. 35, 1862. 
(3 a) Keferstein. — " Beitrage zur anatomiscben und syste- 
matischen Kenntniss der Sipunculiden," in Zeitschrift fiir 
wiss, Zoologie, Bd. xv. p. 404, 1865. 

(4) RouLE. — " Sur le Gephyriens des grands fonds de la mer 

recueillis par le ' Travailleur ' et le ' Talisman,' " in Oomptes 
Rendus, cxxvii. p. 197. 

(5) Selenka. — " Die Sipunculiden," in Semper's ' Reisen.' Wies- 

baden, 1883. 

(6) 0. Ph. Sluiter. — " Beitrage zu der Kenntnis der Gephyreen 

aus dem Malayischen Archipel," in Natuurkundig Tijdschrift 
veor Nederlandsch-Indie, Deel xlv. p. 472, 1886. 

(7) Shipley. — " On Phymosoma varians" in Quart. Journ. Micr. 

Sci. vol. xxxi., n. s. p. 1, 1890. 

(8) Lankester — " Golfingia macintoshii, &c." Trans. Linn. Soc. 

Lond. 2nd ser., Zool. vol. ii. p. 469, 1885. 

Plate VI. 

Fig. 1. The entire animal, magnified about 6^ times. A, the anterior muscular part. 

B, the middle thinner portion. C, the bag-like posterior piece. AN., the 
anus. pp. 29, 30. 

Fig. 2. Semi-diagrammatic figure of a dissection, magnified about 13 diam. I, the 
introvert. E, the e3'e-spots. B.T., the uephridia. H, dorsal vessel. 
D.R., dorsal retractor muscles. V.R., ventral retractors. AN., position of 
anus. Gr, coiled part of gut. F, straight part. pp. 30, 31. 

Fig. 3. Cuticular glands in vertical section. X 290. E.M., radial muscles. 

C, cuticle. L, secretory cells. T, granular contents, p. 30. 
Fig. 4. Hooks from introvert. Surface view. X 154, p. 30. 

Plate VII. 

Fig. 5. Isolated hooks. X 290. F, furrow, p. 30. 

Fig. 6. Transverse section 1 mm. behind the anus. X 53. C.G., cuticular glands. 
R.M., radial musculature. L.M., longitudinal do. M, muscles attached 
to rectum. Gr, rectum. B.T., nephridia. T, tentacle cut across. 
H, hooks. I, introvert. N.C., ventral nerve-cord and muscles, p. 31. 

Fig. 7. Histology of the retractor muscles (Powell & Lealand yV-imm., eyepiece A). 

A. Transverse section of dorsal retractor, showing amount of interstitial 

connective between the bundles. 

B. Ditto of ventral retractor, showing almost no intermuscular connective. 

p. 32. 
Fig. 8. Transverse section of tentacle. X 290. B.S., blood-sinus. N, nerve. 

A, ciliated mesial cells. B, columnar outer cells, p. 32. 
Fig. 9. A. Epithelium of mid-gut. X 290. B. Epithelium of hind-gut. X 290. 

p. 33. 
Pig. 10. Transverse section of oesophagus. X 290. Part only of the stout muscular 

wall is shown. M, muscular wall. E, the epithelium thrown into papillae. 

p. 32. 


Plate VIII. 

Fijj. 11. Transverse section of brain behind origin of ccsoiiliageal cotnTnissures, to 
show triant ganglion-cells. X 290. K.M., retractors. 13.S., blood-sinus. 
G.C., giant cells, p. 33. 
Figs. 12, 13, 14, 15. Four transverse sections through the brain and base of the 
tentacular crown, to show the epineural canal and formation of the cyc- 
spots. X 53. B.S., blood-sinus. V.C., nerve-cord. K.M., retractor muscles. 
S.E., sensory epithelium of the dorsal, and S.V., that of the ventral wall of 
the epineural canal. T, tentacles. S.P., eye-spots. E, epithelium of 
anterior cerebral surface. ]). 31. 
Fig. 16. Transverse section through brain and bases of the tentacles, showing the 
origin of the circuuuesoiihagcal commissures (X 53 and reduced), p. 31. 
A. Circunuesophageal nerves. 

15. Nerves to the epineural canal and dorsal pair of tentacles. 
C. Two nerve-strands, in cross section, which seem to be connected with 

the sense-organs. 
T). Fused retractor muscles. 
K. IJlood-sinus. 
Fig. 17. Diagram of a nephridium. A, excretory canal. B, coclomic pore. 

C, vesicle. D, tubular part. E, body-wall. p. 36. 
Fig. 18. To show histology of tubular portion. X 290. A, peritoneal cells. 
15, muscular wall. C, secreting cells which at D are seen forming 
feathery columns. E, vesicles, p. 36. 

5. On Potamon (Fotamonaufes) latidactylani, a new Fresh- 
water (,'ral) from Upper Guinea. By Dr. J. G. de Man, 
of lerseke, Holland.-^ 

[Received November 15, 1902.] 
(Plate IX.'O 

In the year 1881, Poto.?»o?? o/rica7m?>i A. M.-E. was known only 
by the short diagnosis and the figures in the ' Nouvelles Archives 
du Museum,' ' made from a quite young individual from the 
Gaboon. The anterior legs had neither been described nor figured, 
and it is therefore not surprising that some older specimens of a 
Fotamoit from Liberia were referred l)y me erroneously to this 
species \ 

Some time since, three adult specimens of a Potamon were sent 
me for examination by Prof. Jefirey Bell ; they had been collected 
in the River Prah, in the south of Ashanti, West Africa. These 
Crabs not only proved to belong to the same species as that 
described by me in 1881, when compared with a female of medium 
size and a veiy young male from Libei-ia in the Leyden Museum, 
but they proved also to be new, as a typical specimen of P. afri- 
caimm, a middle-sized female from " Ogoue " (evidently the 
River Ogowe, just below the Equator), was khidly sent me by 
Prof. Bouvier, and as a more complete description of P. africanum 
was published in 1887, in which, however, the legs have not licun 

1 Communicated by F. Jeffeey Bell, F.Z.8. 

- For explanation of the Plate, see p. 4-7. 

3 Vol. V. p. 186, pi. xi. fig. 2 (1869). 

* Telpkusa africana de Man, Notes Leyd. Mas. iii. p. 121 (1881). 

42 DR. J. G. DE MAN ON A NEW [Jan. 20, 

figured (A. Milne-Edwards, ' Observations stir les Crabes des 
eaux douces de TAfrique/ Paris, 1887, p. 4, pi. 2. fig. 8). I 
therefore propose the name of Potamon latidactylimi for this new 
species that inhabits Liberia and Ashanti, on account of the 
characteristic shape of the hands, and as this feature was not alluded 
to in my somewhat incomplete description of 1881, another will, 
I think, be welcome. 

The rivers of West Africa and of the Soudan are inhabited by 
several species of PotaTiion which differ from the other species 
of this subgenus in the existence of tioo epibranchial teeth behind 
the external orbital angle, instead of one as is usual. I was 
at first inclined to create for these species a new subgenus, but 
the difi'erences from Parathelphusa are, indeed, of too little 
importance. These species are the following : — 

Potamon auhryi H. M.-E. Gaboon. 

,, peJii Herklots. Gold Coast. 

,, africanum A. M.-E. Gaboon ; French Congo. 

,, decazei A. M.-E. French Congo. 

,, emarginatibm Kingsley. West Africa. 

,, floweri de M. Soudan. 

,, latidactyltim, n. sp. Tipper Guinea. 

It must, however, be observed that P. emarginatum is con- 
sidered by Miss Rathbun (Proc. U. S. National Museum, xxii. 
1900, p. 283) to be the same as P. auhryi. P. latidactylum 
may at first sight be distinguished from P. africanuvi hy the 
different shape of the extraorhital and epibranchial teeth and hy 
the different form of the hands, especially of the fingers, but there 
are still other difi'erences. 

The cephalothorax is a little more enlarged than in the species 
from the " Ogoue," as is readily shown by the measurements. 
The upper surface is somewhat less depressed, the branchial regions 
are somewhat swollen, especially the anterior ones, and the gastric 
and cardiac areas are, in aged individuals, also slightly convex and 
not so much depressed as in P. africanum. The oblique furrows 
or depressions limiting off" the protogastric areas from the anterior 
branchial lobes are scarcely distinguishable in P. africamim ; in 
the other species, however, they are quite distinct, and, in adult 
individuals, rather deep. The urogastric lobes and the cardiac 
area are a little hroader in proportion to the breadth of the 
carapace than in P. africanum. In the latter the gastric region 
is faintly granular or rugose just behind the postfrontal ridge, 
but in the new species it appears everywhere smooth, though 
finely and sparsely punctate. The anterior branchial area is 
slightly rugose in P. africanum., but not marked with oblique 
wrinkles ; these are quite distinct in the species from the River 
Prah and Liberia ; on the posterior branchial lobe these rugosities 
occur in both species, they are, however, thinner and finer in 
P. latidactylum. The posterior part of the upper surface 

PZ.S. 19 03, vol. I. PL IX. 


y IMS 



J. G.deMan del. 


^-I 'OLJ 


appears, on either side of the cardiac and intestinal regions, 
somewhat granular and rugose in the Congo species, but^'^^e 
smooth, though finely punctate, in P. laiidactyliim. 

In the large aged female from the River Prah the oblique ruga3 
near the lateral boundaries of the carapace show a tendency "to 

In both species the postfrontal ridge is rather prominent, 
smooth, and nowhere granulated. In the type specimen of 
P. africmmm it extends in a nearly straight hne toAvards the 
second epibranchial tooth, uniting with that of the right side, 
but ending about 1 mm. short of that on the left. The postfrontal 
ridge of P. latidactylum usually curves slightly forwards mesially 
and at each end ; it never unites with the epibranchial teeth, 
ending about 1^ or 2 mm. short of the middle of the first epi- 
branchial tooth. It is situated a little more distant from the 
orbits than in P. africanum., so that the furrow behind the orbits 
is somewhat narrower in the latter species. The front has the 
same form and breadth in both, and shows in the middle a broad, 
though shalloio hay ; both the front and the furrow behind the 
orbits are a little granular in P, africantmi, but quite smooth, 
though finely and sparsely punctate, in P. latidactylum.. 

The extraorbital and the two epibranchial teeth have a qidte 
different form (PI. IX. figs. 1-3 & 7, 8). In P. africammn 
(PI. IX. figs. 7, 8) they are more jwominent and separated from 
one another hy much deeper incisions. The first epibranchial 
tooth is, in this species, a little smaller than the extraorbital, but 
has about the same form ; its straight or slightly arcuate outer 
margin makes a right angle with the anterior margin of this 
tooth. In P. latidactylum, however, the extraorbital tooth and 
the anterior epibranchial one are much less prominent and mttch 
loiver, the anterior margin of the first epibranchial tooth being 
considerably shorter than its outer margin, measuring only one-fifth 
of it. In the aged female from the Eiver Prah the extraorbital 
tooth appears somewhat longer than the epibranchial one, and in 
the two males (PI. IX. fig. 1) they have about the same length; 
in the much younger female from Liberia (PL IX. fig. 3) the 
epibranchial tooth, however, is almost twice as long as the extra- 
oi-bital, but in the quite young male from the same locality 
(PI. IX. fig. 2) the extraorbital tooth appears again once and a 
half as long as the epibranchial. As regards the relative length 
of these teeth, P. latidactylum presents therefore considerable 
individual difierences. In both species the second epibranchial 
tooth passes backward into an arcuate crest, bounding the cephalo- 
thorax later;il]y ; this crest, which in young individuals of P. lati- 
dactylum (PI. IX. fig. 3) appears distinctly denticulate, the den- 
ticulations or crenulations disappearing in more aged individuals, 
reaches in P. africanum to th^ 2^osterior boundary of the tiro- 
gastric lobes and does not curve imoard on to the surface of the 
branchial region : in P. latidactylum, however, it is much shorter, 

44 BR. J. G. DB MAN ON A NEW [Jnil, 20, 

scarcely reaching to the level of the mesial crescentic portion of 
the cervical suture, and it distinctly curves inwards on to the 
upper surface of the carapace. The orbits have the same form. 
in both species, but in P. africanuin the incision that separates 
the extraorbital tooth from their lower margin is deep, much 
deeper than in the new species, so that in the former the outer 
angle of the orbits strongly projects beyond this margin. 

In P. africanum the transverse furrow, limiting off the sub- 
hepatic region from the branchial floor, is bordered by a row of 
small granules, the subhepatic area is covered with prominent 
rugosities and granules ; in our new species the latter appears 
almost smooth, and the row of granules is also less distinct. The 
rugosities on the inflected portion of the cephalothorax and on 
the outer part of the pterygostomian regions are also much less 
distinct in the species from Upper Gviinea than in P. africanitm. 

The outer foot-jaws have the same form and charactei's in both 
species, and the furrow on the ischium-joint runs in both a little 
closer to the internal than to the outei- margin of this joint. 

The type of P. afo^icanimi is a female, the abdomen of which 
has not yet obtained its full development and size. The sterniim 
of P. latidactylum is smooth, punctate, and in both species a 
transverse furrow unites the postero -external angles of the buccal 
frame. The male abdomen (PI. IX. fig. 4) somewhat resembles 
that of P. consohrinum de M. (Notes Leyden Musevim, 1899, 
pi. 10. fig. 10,?"). The lateral margins are slightly concave. The 
terminal joint has about the same length as the penultimate ; it 
is triangvilar, rather pointed at the tip, not rounded like that of 
P. consohrinum ; the lateral margins are veiy slightly arcuate 
posteriorly, nearly straight towards the tip. The posterior margin 
of the penultimate joint is a little broader than the anterior ; the 
lateral margins, slightly convex anteriorly, are a little concave 
posteriorly. The length of the antepenultimate joint measures 
about two-thirds that of the penultimate, its posterior margin is 
a little concave, so that the lateral margins are somewhat shorter 
than the length of this joint in the middle line. The lateral 
margins of this and of the following fourth joint are sti'aight, but 
those of the fifth are rounded ; in P. consoh-inum, however, this 
latter is also straight. As regards the coarse punctation, P. lati- 
dactylum agrees with P. consohrinum. 

In the two males from the River Prah the right chelipede is a 
little larger than the left ; in the very young one from Liberia 
the left is a little larger. The upper margin of the arm of the 
larger male from the River Prah bears some transverse i-ugosities, 
and one observes a small tubercle on the concave upper surface at 
the proximal end, somewhat nearer to the upper than to the 
anterior margin ; the latter is also somewhat tubercular on its 
proximal half. The anterior surface of the arm bears, near the 
articulation of the wrist, a conical tubercle not far from the 
anterior margin, and a lower, broader tubercle between it anr' 


the under margin; the latter is also sh'ghtly tubercular alonc' 
its distal lialf. The three faces of the arm are quite smootli. 
The carpus is also nearly smooth above, though spai-sely and 
rather finely punctate ; the inner margin is armed witli a i-ither 
small pointe<l spine in the middle and beneath it with a small 
acute tubercle. The larger chela (PI. IX. fig. 5), measured 
horizontally, is a little longer than the length of the ce})halotliorax 
and the fingers are a little longer than the palm ; the latter is 
just as long as high near the ai-ticulation of the fingei-s. The 
convex outer suiface of the palm is quite smooth, though finely 
punctate, similar to the upper margin, but the lower edge of the 
palm is ol)soletely tubercular. The fingei-s are someichat com-pressed 
especialh/ the rmmobile one. I'he immobile finger is ratlier hiqh 
at the base; \ti^ flattened outer surface appears minutely and 
densely granular under an ordinaiy lens, and one observes on it 
three longitudinal rows of impressed points running to the tip of 
the finger, the middle one of which is situated a little closex- to 
the lower margin of the finger than to the ujipei-, and looks 
like a very shallow furrow. The 15 or 16 teeth are small, two 
or three are somewhat hu'ger than the others, and one in the 
n\id<lle of the finger is the lai-gest of all. The fingers leave a 
narrow interspace between them when closed, the pointed tips 
being only in contact and crossing one another. The dactvlus is 
somewhat arcuate and tapers rather slowly to the tip ; both on 
the outer and inner sui'faces and on its upper edge this finger 
appears minutely gramdar under a lens, and one observes on it 
several rows of small shallow puncta. The teeth agree in size and 
in number with those of the index, three or four being somewhat 
larger than the others. The little convex inner surface of the 
palm is quite smooth. 

The left chelipede is somewhat smaller, the hand beino- 41 mm. 
long ; it fully agrees with the right one, but the fingers are in 
contact and the I'ows of puncta ai'e a little moi-e distinct. 

In the young male from Liberia the fingers are in contact in 
both chela?, the upper margin of the palm appears distinctly 
granulate under a lens, and the longitudinal furrows on the 
fingers are already visible to the naked eye. 

In the young female from Liberia the right chelipede is a little 
larger than the left, the fingers are in contact, and the furrows 
on the index, as also the rows of puncta on the mobile finger, 
are distinctly visil)le to the naked eye. The upper margin of the 
palm of the left hand appears finely gramdar. 

In the aged female from the River Prah the left chelii^ede is 
much larger than the right ; the hand, indeed, is 45 mm. long, 
just as long as the cephalothorax, and 20,| mm. high, but the 
right chela (PI. IX. fig. 6) is only 31 mm. long and 13 mm. high. 
In both the fingers are in contact throughout their whole len<>1:h 
and the pointed tips cross one another ; the strongly compressed 
and much flattened immobile finger is not furrowed, though the 

46 DR. J. G. DE MAN ON A NEW [Jan. 20, 

rows of puncta are still visible. The fingers are but little longer 
than the palm, and the latter is on its outer surface and on its 
upper margin smooth, sparsely punctate. The fingers of the 
smaller chela, however, are once and a half a,ii long as the palm 
(PI. IX. fig. 6). 

The female type-specimen of P. africanum compared with the 
female from Liberia, which is about the same size, shows the 
following differences : — The anterior legs (PI. IX. fig. 9) are equal, 
as^ regards their shape and size. The transverse rugosities on 
the upper margin of the arm are more distinctly granular, and 
the anterior and lower margins are also more distinctly tuber- 
culated ; the conical spine on the anterior surface, near the articu- 
lation of the wrist, is more pointed, and several small tubercles 
are seen between it and the anterior margin of the arm, that are 
wanting in P. latidactylum. The upper surface of the carpus is 
slightly granular at the base of the spine on the inner margin, 
and one observes, moreover, everywhere fine impressed lines and 
points ; the spine is somewhat larger than in P. latidactylum, 
and instead of a small tubercle, there is beneath it a smaller, 
pointed spine. The hands (PI. IX. fig. 9) have a quite different 
form. The fingers, almost in contact, are distinctly somev:hat longer 
than the palm, and the latter is distinctly less high than long. The 
upper margin of the palm is covered with depressed granules, 
and the puncta on the convex outer surface are partly arranged 
in longitudinal rows. The fingers are not compressed, slender, 
7ior curved, except at the pointed tips which cross one another. 
The immobile finger is much loioer at its base than in the 
other species, and its convex outer surface is distinctly furroived ; 
of the three furrows visible to the naked eye, the middle one is 
deeper than the others. This finger is armed with 25 or 26 small 
teeth, some of them on the pi'oximal half are somewhat larger 
than the others. Finally, the dactylus tapers regularly to the tip 
and is also longitudinally furrowed by rows of impressed points ; 
the teeth agree with those of the index. The fingers are almost 

The ambulatory legs, the measurements of which are giA^en 
on p. 47, apparently agree with those of P. latidactylum, but the 
meropodites are -inore granular along and near their upper edges. 

Potamon auhryi H. M.-E., from the Gaboon, P. pelii Herklots, 
from the Gold Coast, and P.floweri de M., from the Soudan, are 
different species (mcZe de Man, Proc. Zool. Soc. Lond. 1901, vol. i. 
p. 94), being at first sight distinguished by the upper surface of 
their cephalothorax being very convex from behind forwards, 
smooth and shining, hy the different form of the fingers, of the 
abdomen, &c. The young male of Potamon pelii described I. c. 
p. 99, I have re-examined for the purposes of this memoir. 


Jleasurements in millimetres. 















35. V 



















































































Breadth of the cephalothorax 

Length ot'tlie cephalothorax without the uhdomen... 

Distiince between the external ovliitiil angles 

„ „ posterior eiiihraiichial teeth ... 

Breadth of tho anterior margin of the front 

Distance, in tho niwlian line, between the anterior 

margin of the front and the postfrontal ridge 

Distance between the outer angle of the orbits and 

t he post frontal ridge 

Distance between the outer angle of the orbits and 

t he posterior epibranchial tooth 

Breadth of the iirogastric arciu 

„ „ cardiac areai 

,, ,, orbits 

Height of the orbits, near the lower internal angles . 
Length of the terminal segment of the abdomen 

,. ,, penultimate segment 

Breadth of the anterior margin of this segment 

,, ,, posterior margin of it 

Length of the larger chela 

„ „ palm 

Height of the palm, near the articulation of the 


Length of the legs of the penultimate pair 

„ „ meropodites of the penultimate pair... 

Breadth „ 

Length of the propoditcs „ „ „ ... 

Breadth „ „ „ „ „ ... 

Length of the dactylopodites ,, „ „ ... 

„ „ legs of the last pair 

„ „ meropodites of the last pair 

Breadth „ „ ,, „ 

Length of the propodites „ „ 

Breadth „ „ „ „ 

Length of the dactylopodites „ „ 

48i 34J 
33 24| 

my. 23 




















22i 34 
11 116 






6i . 
91 15.V 
4 Q\ 

111 j 18 

39 60 




Nos. 1-3, River Prah ; Nos. 4, 5, Liberia; No. 6, type-specimen of Theljiihusa 
africana A. M.-E., Ogouc, from the I'aris Natural History Museum. 


Figs. 1-6. Potamoii (Potamnnautes) latidacti/him, n. sp. — Fig. 1. Dorsal view of 
the anterior part of the carapace of the largest male specimen from tho 
River Prah, X U-. Fig. 2. Left antero-external angle of the carapace of 
the young male from Liberia, X 3. Fig. 3. The same of the young female 
from Liberia, X 3. Fig. 4. Abdomen of the largest male from the River 
Prah, X 11. Fig. 5. Larger chela of the same male from the River Prah, 
X 11. Fig. 6. Smaller chela and carpus of the adult female from the 
River Prah, X 11. 

Figs. 7-9. Potamon (Potamonautes) africamim A. M.-E., female type-specimen of 
the Paris Jfuscum, from Ogoue.— Fig. 7. Dorsal view of the anterior part 
of the cephalothorax, X 11. Fig. 8. Left antero-external an-le of the 
carapace, X 3. Fig. 9. Right chclipcde, X l-V. 


6. On a new " Bird's-dung ^' Spicier from Ceylon. By R. I. 
PococK, F.Z.S., and the Hon. N. C. Rothschild, B.A., 
RE.S., F.Z.S. 

[Received November 18, 1902.] 
(Plate X.^) 

During a recent visit to Ceylon, the Hon. N. C. Rothschild, in 
company with Mr. E. E. Green, discovered a specimen of a 
Spider (Phrynarachne) imitating, for purposes of allvirement and 
concealment, a patch of bird's-dung. A photograph of the spider 
resting upon the leaf iii situ was obtained, and the leaf with its 
patch of web was brought home with the spider (PL X. figs. 1, 2), 

The film of white web upon the leaf, with its irregular outline 
and marginal, often apically thickened projections, closely assimi- 
lates the splash of the fluid components of the excrement ; while 
the spider itself, with its mottling of black and white, resting 
upon the centre of the film, with its legs tucked in, represents 
the semi -solid central core. So close, indeed, was the imitation 
that both observers were for a time deceived, until closer scrutiny 
revealed the true nature of the deceptive patch. 

It is commonly assumed that the purpose of this imitation is 
purely alluring or pseudepisematic. ISTo doubt, however, it is an 
equally impox'tant pi'ocryptic factor, serving to protect the spider 
from various enemies, especially the wasps of the family Pompilidse. 

The spider, which appears to represent an undescribed species, 
may be named after one of its discoverers ^. 

Phrynarachne rothschildi, sp. n. 

2 . — Colour. Carapace with a broad central black patch, with 
irregular lateral margin, extending from the eyes to the posterior 
slope, and forming an angular expansion on each side imme- 
diately behind the head ; and with its posterior border passing 
into a median longitudinal stripe, continuous with a trans- 
versely arcuate black stripe which extends from the middle of 
the posterior slope of the carapace on to its lateral portion, 
breaking up into fainter patches separated by radiating pale 
bands ; clypeus, sides of head, and area between central patch and 
lateral patches yellowish white. Mandibles yellow in basal half, 
with a large black patch, not reaching the fang, in the distal half 
in front, blackish brown benea,th. Sternum mostly yellowish white, 
with a large subtriangular black patch, di-\dded by a narrow pale 
line in its posterior half, and some black at the bases of the legs. 
Labium blackish, maxillfe blackish internally, whitish externally. 
Palp with its femur white below, black above in its basal half ; 
patella white, tibia and tarsus black, variegated with yellow 
patches. First and second leg's with the coxae black, variegated 

1 For explanation of the Plate, see p. 51. 

2 Mr. R. I. Pocock takes the sole responsibility for tlie naming and description of 
the species. 

p. Z. S. 19 03^ vol 1. PI X 

F.Hckard- Cambridge del eftlith, 


West.Nevnnan imp. 

1001).] A Xi:w SPIDKU KIIOM CEYLOX, 49 

with dirty yellow below, whitish witli a black spot above ; femora 
jet-black variegated with paler patches alx)ve, in front, and 
behind, with a uairow whitish line between the tubercles below, 
the distal end yellowish white; patellar yellowish white, with 
two palely fuscous patches above and some behind ; tibise and 
protarsi mostly jet-black, with a, pale median inferior longitudinal 
stripe between the spines; tarsi pale above, black below; third leg 
with coxa and ti-oehanter blackish below, the femur amber-yellow 
in its liasal two-thirds, yellowish white with a black [)atch, much 
broader liehind than in front, in its distal third ; tibia black and 
whitish, the l)lack predominating posteriorly; protarsvis and tarsus 
mostly yellowisli; fourth leg, with exception of its coxa, mostly 
yellowish whit(% lightly spotted above. Almost the whole of the 
upperside of the abdomen jet-black from its anterior margin 
backwards to the spinners, with the postero -lateral tubercles and 
the sides yellow ; the under surface {i. e. the area between the 
lung-sacs on the epigastric region and behind the genital fold 
nearly back to the spinners) black, this black fielil showing a ti'ans- 
vei'sely oblong pale patch behind the genital apeiture (Plate X. 
figs. 3 & 4). 

Carapace not strongly tubercular, flattish longitudinally along 
the middle line, with a shallow transvei'se depression behind the 
ocular area of the head ; area between the ocular tubei-cles 
depressed ; eyes of posterior line (fig. 5) subecpially spaced, the 
medians about 5 diameters apart, lying about their own diameter 
in front of a tangent joining the anterior borders of the lateials, 
which are distinctly largei- than the medians ; eyes of anterior line 
(fig. 6) strongly recurved, the inferior edges of the latei'als as high 
as the superior edges of the medians, the latter considerably smaller 
than the laterals, about 2.| diameters from each other and about 
2 diameters from the laterals ' ; distance between the medians 
about equal to the height of either above the edge of the clypeus, 
which is weakly tiibercular and furnished with ^n'orainent angles. 

Mandihles prominent basally, scarcely tubercular ; margins of 
fang-groove thickly fringed, the posterior aimed with one fang, the 
anterior with teeth, remote from base of fang; the posterior fringe 
continuous with a scanty band of hair passing up the inner side 
of the posterior surface of the mandible. 

Legs. Femur of 1st leg thickly covered in front with small 
tubercles, with a few large tubercles above and two rows of about 
five tubercles each below, the posterior surface smooth ; tibia 
bowed, armed beneath with two rows of about 4-5 spines, the 
anterior and dorsal sin-face also spined ; protarsus a little longer 
than tibia, armed beneath on each side of the middle line with a 
band of longer and shorter inegularly arranged spines, those on 

' Owinp to ;i want of dotinitinn of tlie bordov of the conipal Ipns, fhi^ pvocise ';i;!e 
of the eyes of the antprior line, mid consequently of the width of the intcrsjiiice 
between them, is hind to iisrertnin, varying: apparently in difli'vint lights and with 
Ifnses of ditterent power. In the ahove-priven description the eyes are described as 
seen under a 4-inch objective and a platyscopic lens. 

Proc. Zool. Soc. -1903, Vol. I. Xo. IV. 4 


the posterior side being much less numerons than on the anterior, 
the anteiior and dorsal surfaces of the segment also spiny ; 
tarsus covered with rows' of spinules ; 2nd leg like the 1st, but 
with the femur almost quite smooth ; 3i'd and 4th legs with tarsi 
bristly beneath, tibia of 3rd spined below, of 4th scarcely spined. 

Abdomen as wide behind as it is long, and twice as wide as it is 
in front, the anterior border ti'ansvei'se and armed with four 
tubercles, the lateral border with four small tubercles, the 
postero-lateral angle with two large superior and two small 
inferior pale tubercles ; between the former are four pairs of 
shining black tubercles, tAvo pairs of which are mvich larger than 
the others ; in addition to these the dorsal surface is furnished 
with three pairs of small scattered tubercles ; lateral surface 
pitted ; inferior surface with two rows of muscular pits. Vtdva 

Measurements in mm. — Total length 9, carapace 4 ; posterior 
width of abdomen 5'5, anterior width nearly 3 ; length of 1st leg 
14, its femm^ 4, patella + tibia about 5, 3rd leg 6, tarstxs + pro- 
tarsus about 5. 

Log. Kandy. 

In size and shape this species does not appear to differ appreci- 
ably from F. ceylonica 0. P. Cambridge (P. Z. S. 1884, p. 201, 
pi. XV. fig. 3, sub Ornithoscatoides), which was recorded from 
Ceylon. The colouring of the two, however, is very different. 
In F. ceylonica the anterior legs are heavily blackened only upon 
the protarsi and distal half of the tibige, the mandibles have no 
black patch, and the black-and-yellow pattern of the carapace 
and abdomen, so conspicuous in P. rothschildi, is but little differ- 
entiated ; also the cephalic tubercles between the two rows of 
eyes are much higher in F. ceylonica (see also Simon, Hist. Nat. 
Araign. i. p. 1043, fig. 1087, 1895). 

Another Ceylonese species, 0. nigra 0. P. Cambr. (Joe. cit. 
p. 202, pi. XV. fig. 4), was based upon the male sex, and cannot, 
therefore, be compared with those based upon females. A third 
species, F. fatalis O. P. Cambr. (P.Z. S. 1899, p. 525, pi. xxx. 
fig. 7), from Ceylon, differs so markedly in form, coloration, 
tuberculation, &c., as to need no comparison with this new species. 
Also F. peeliana Stol. (J. A. S. Bengal, xxxviii. 1869, p. 229, 
pi. XX. fig. 4), from Sibsagar, Assam ; F. Uiberosa Blckw. (Ann. 
Mag. N. Hist. (3) xiv. p. 38, 1864, & P. Z. S. 1884, pi. xv. fig. 2), 
from the East Indies ; and the Burmese foi'm F.papiolaia Thorell, 
are all different from this new form. 

To Dr. H. 0. Forbes, F.Z.S.', belongs the credit of the discovery 
that the coloration of the species of Fhrynarachne belongs to the 
pseudepisematic category. The pattern of yellow and black which 
decorates the dorsal and ventral sides simulates that of the semi- 
solid central portion of a pa,tch of biixl's-dung splashed upon a leaf, 
the paler more fluid portion being represented by a thin irregular- 
shaped carpet of white silk, in the centre of which the spider 
takes its stand. The spider discovered by Messrs. Ptothschild 
1 P. Z. S. 1883, p, 586, pi. li. 

1903.] A XEW sriDE!i rnoM ckylox. 51 

and Green was resting upon the leaf in tlie uunnal position, tliat 
is to say back uppermost (figs. 1 &> 2). 

Forbes, however, expressly states that F. decipiens, the species 
he discovered, lies back downwards on the wel), holding itself 
in place by means of the sjiines with whicli the anterior upper 
surfaces of the legs are furnisheil, and he adds that the under- 
side of " its rather irregidarl y-shaped and prominent aljdomen 
is almost all white, of a pure chalky white" (p. 587), and that 
" its piu-e white abdomen represents the central mass of the bird's 
excreta, the black legs the dark poi'tion of the slime." Refer- 
ence, howevei', to his figure, which was apparently drawn partly 
from memory, the leaf and web having " gone astray " during 
the transport home, shows that the iniderside of the abdomen, so 
far from being almost all white, is furnished with a large black 
sub-oblong patch, which extends from its anterior boi'der con- 
siderably past the middle. It is noticeable, too, that from the 
ventral aspect the abdomen is not iiTegularly shaped, but evenly 
oval in outline and devoid of tubercular excrescences. The dorsal 
surface, on the contraiy, as repi'esented in the figui-e given by 
INIr. Cambridge (P. Z. S. 1884, pi. xv. fig. 1), might very well 
be described as almost all white and rather irregularly shaped. 
The white colour largely predominates, and the tubercles project 
prominently fi-om the expanded posterior portion. ISIention is 
made of these discrepancies between the figure and description, 
to justify the suspicion that Di*. Forbes may have mistaken the 
dorsal for the ventral surface of the spiders he saw. Moreover, 
one cannot but wonder how the spider maintains a secure hold 
back downwards, especially when the powerful prehensorial legs of 
the first and second pairs are released, as released they m\ist 
surely be, to seize an alighting butterfly. One would think that 
the flapping of the insect's wing would pull the spider, now 
insecui-ely anchored, from its hold and bring both to the ground 
together. On the other hand, if the spiders of this kind always 
rest in the normal position seen and photographed in the case of 
F. roihschildi, the simulation of the bii'd's-di;ng is equally perfect, 
and the spider, while seizing a butterfly with its fore legs, can 
maintain itself securely in place by gi'asping the web with the 
claws of the remaining pairs and by gluing its spinning-mamilke 
to the subjacent silken threads. 

These, then, ai-e the points for further investigation which we 
would ask i-esidents in the tropics to take note of : — Do the species 
of Fhrynarachne assume indifl:erently the dorsal or the ventral 
attitude ? Is it a peculiarity of one species to lie back uppermost, 
and of another belly uppermost ? If they lie, as Forbes asserts, 
back downwards, how is a secure hold maintained when an insect 
has to be seized ? 


Fhrynaraclviie rothscJiildi, sp. n., p. 48. 

Figs. 1, 2. Tlie Siiiilcr vesting on leaf. Fijr. 3. Up])er suiiacc. 

Fi^-. i. Under surface. Fics. .i, 6. Eves. 



7. On the Crustaceans of the Genera Petalidhim and Sergestes 
from the ' Challenger/ with an Account o£ Luminous 
Organs in Sergestes cliallengeri, n. sp. By Dr. H. J. 
Hansen (Copenhagen) . 

[Received November 29, 1902.] 
(Plates XI. & XII.') 

Dui'ing a stay in London in July and August, 1902, I examined 
various groups of Crustacea in the British Museum (Natural 
History). I beg the Director, Professor E. Ray Lankester, 
and Mr. F. Jeffrey Bell to accept my sincere thanks for the free 
use of the collection and for their kind help. 

In the paper " On the Development and the Species of the 
Crustaceans of the Genus Sergestes " (Proc. Zool. Soc. Lond. 
1896, pp. 936-70) I have given a revision of this extensive 
genus. I had studied a very rich material of pelagic forms 
belonging to the Copenhagen Museum, among which are all the 
types of Kroyer ; besides I had examined types of 5 species 
established by Chun, Metzger, and Ortmann. 

Among other things, I proved that " of the 59 {or 60) hitherto 
described species only about 20, or one-third of the total number, 
have been established on adult animals, such as have almost or 
entirely arrived at sexual maturity ; and that almost all the other 
species are true larvce, and even of these a considerable mmiber are 
larval stages of species already established on adult specimens, . . ." 
Of earliei- authoi-s, C. Spence Bate has produced a very large con- 
tribution on the genus Sergestes, extending to eighty-eight quarto 
pages and seventeen plates, in his " Report on the ' Challenger ' 
Macrura." He established the genus Petalidium on a new species, 
described 24 new species of Sergestes &c. In 1896 I wrote 
(p. 939) : " This large contribution is of course of great 
importance, but unfoi'tunately neither the descriptions nor the 
figures are so good as could be wished, and in numerous instances 
... a re-examination of the type specimens is absolutely necessary 
— the greater part of the new species are but larvas." I have now 
studied all types which are preserved in the British Museum, and 
the present paper contains the results of my examination. 

Bate describes 31 species of Sergestes as examined by himself : 
of these, 24 are established as new to science, 6 are considei-ed 
to be Kroyerian species, and one is referred to S. atlanticus 
H. M.-Edw. The types of 9 of the species established by Bate do 
not exist in the British Museum ; some specimens mentioned in 
his work and belonging to other species are also absent ; but 
several specimens belonging to varioiis species and omitted in the 
Report were found in the collection. I am therefore only able 
to give more or less incomplete notes, based on the study of the 

1 For explanation of the Plates, see p. 78. 

PZ S 1903, vol. I. PI, XI 

H. J. Han. 

j.y Edwir. Wilson , Camhnt/ae 


P.ZS. 1903, vol. I PI. XII 

del. Edwin Wilson. Cambridge 



specimens, on 22 of the species in question, 15 of which were 
established by Bate himself. Furthermore, he refers Petalidium 
Bate, Sciacaris Bate, Acetes H. M.-Edw., and Lucifer Vaugh. 
Tliomps. to his family Sergestidte ; of Lucifer he describes two 
species, but the genus must be more thoroughly studied than has 
hitherto been done before the examination of the ' Challenger ' 
specimens ; Sciacaris Bate has one species, which is only a larva of 
a Sergestes, and the tyjje seemed to be wanting in the Museum ; 
of Acetes, Bate had no specimens; and Fetalidium is mentioned 

Of Bate's 31 species of Sergestes only 6 arc really mature forms, 
25 being larvie. Special attention has been jjaid to the adult speci- 
mens j)i'esei'ved in the Museum and enumei-ated by Bate ; on two 
of these specimens I have established two new species, and 
besides I add notes and some drawings to the representations of 
Bate. Unfortunately nearly all the specimens of rare and 
especially interesting species were veiy much mutilated. 

Our pi'esent knowledge of the adult species of Sergestes of the 
Atlantic fauna is far from complete, liut yet we ai'e acquainted 
with so many species that it was possible for me in my earlier 
paper either to refer the Mastigopus-iovYL\^s, examined to the 
mature species, or to describe the older larval stages and some- 
times the black-eyed but still immature forms, so that they can be 
recognized with certainty and referred to the mature forms when 
these are discovered in the future. All the Atlantic lai-val forms 
from the ' Challenger ' seen by me have now been referred in a 
similar way. But many lai-vse established by Bate as valid species 
of adult or sub-adult animals have been secured in the Pacific. 
Our knowledge of the mature stage of the species living in that 
vast ocean is still rudimentary ; and I have therefore not been 
able to refer the lai-vfe of three of Bate's species to any species 
established on adult specimens. Bate's types of his species 
established on larval forms are often either defective or very 
young, wherefore I thought it of little use to describe and figure 
them again ; but I have generally added some notes on their 
affinities, and sometimes also a few corrections to his descrip- 
tions. "When the Pacific has been modei'ately well explored 
by further expeditions, many adult forms and their larval stages 
will be discovered ; and a future student of the group will then be 
able to refer at least some of the larva?, which I cannot interpret, 
to their adult foi-ms. To the young larvse described b}' Bate as 
species of Mastigopus I pay no attention at all : the types seem 
to be lost. 

I think it convenient first to deal with all the ' Challenger ' 
species in the Siime consecutive order in which they are described 
in Bate's Report ; then to put together some residts of the investi- 
gation ; finally, to mention more fully tlie luminous organs in 
Sergestes chcdlengeri, n. sp. 

In order to abridge the descriptions, in the following pages — 
as in my earlier paper^ — I make use of some abbreviations : — 


antenn. ped, = peduncle of the antennulse ; mxp.^-mxp.^ = the 
second and third pair of maxillipeds ; trl.^-trl.° = the first to the 
fifth pair of trunk-legs ; br. = the first, br.^ = the second branchia 
above the same legs ; ext. br. of urp. = external biunch of the 

I. Notes on the Species of Petalidium and Sergestes. 

a. Petalidium Bate. 

To this genus Bate has referred only one species. It will be 
convenient to deal with the characters of the genus together with 
those of the species. 

Pet. foliaceum Bate, pp. 348-50, pi. Ix. (Plate XI. figs, la-1 g.) 

Bate mentions five specimens from two stations : Stat. 146, 
lat. 46° 46' S., long. 45° 31' E., 1375 fathoms; and Stat. 159, lat. 
47° 25' S., long. 130° 22' E., 2150 fathoms. All these specimens 
are at present in the Museum, but even Bate's text and his figure 
of the entire animal show that most of the appendages presenting 
specific characters are wanting or have been mutilated. Bate 
writes (p. 349) : " The great distinction between this genus and 
Sergestes exists in the form, character, and arrangement of the 
branchial plumes, which consist of a series of plates and cylin- 
drical filaments, situated side by side in a series of rows at right 
angles to the stalk. There is but one plume to each of the five 
anterior somites of the j)ereion, the posterior two somites having 
none ; between some of the somites is a large foliaceous plate." 
But this description is difiicult to understand : his figure 3, 
representing the branchiae, is defective, and his tabular view 
(p. 349) is wrong. He was of opinion that the foliaceous plates — 
of which he had seen only three — were pleurobi'anchise, Avhile the 
"branchial plumes " were arthrobranchiae ; but this is incorrect: 
they are decidedly pleiii-obranchiae as in Sergestes. Petal. foUaceimi 
Bate dififers from all species of Sergestes in one quite unimportant 
feature, that no trace of branchiae is found above trl.'*, and in one 
important character, viz., the sti'ucture of the pleurobianchial 
plumes. This structure is very interesting (fig. 1 e). The most 
developed branchiae are, as usual, those above ti'l.^ and .trl.'^ ; each 
of these consists of an anterior and a posterior half, and each half 
of five (to six) transverse rows of branchial plates, generally five 
or six in each row, and these plates (some of the lowest excepted) 
are directed upwards. The pleurobranchiae above mxp.^ and 
especially trl.^ are less developed, with a lower number of trans- 
verse rows, and partially with a loAver number of plates in the 
rows. The pleurobi'anchial plumes in Fetalidium look very 
difiei'ent from those in Sergestes ; the real differences are : a 
much loAver number of rows, a much lower number of plates in 
the rows, and that the plates are much larger, curved upwards, and 
look much more independent. Behind the upper part of each of 
these four pleurobranchise originates a pleui'obranchial lameUa (Z.), 


which is a I'educed branchia ; these lamellae are very long above 
trl.^ and ti*l.^, while the two anterior lamellse above mxp.^ and 
trl.^ are much less prominent. On mxp." is found an epipod (e/?.), 
with a branchia consisting of a few plates, and above it a rudi- 
mentaiy pleurobianchia consisting of one very small lamella. 

Fig. 1 a and fig. 1 c show the rostrum, consisting of a carina 
with a short or veiy shoi-t terminal spiniform pi'ocess. There are 
no supraocular or hepatic spines ; the gasti'o- hepatic groove is 
well developed. A comparison of fig. 1 c with fig. 1 d shows that 
the eye-stalks are considerably depi'essed ; they are from two and 
a half to neai-ly three times longer than the eje& at the inner 
terminal angle, with a small rounded knot turning inwards and 
a little upwards (fig. 1 h ; fig. 1 d), and besides (always ?) with an 
exceedingly small tubercle somewhat in front of the inner angle 
and more downwards on the inner side (fig. 1 d). In the antenn. 
ped. the basal joint is very broad (fig. 1 &), decidedly shorter than 
the outer margin of the two other joints together ; the third joint 
is scarcely three times longer than broad, considerably longer 
than the outer margin and a little shorter than the inner mai-gin 
of the second joint, which is stout, with its inner margin only a 
little moi'e than twice as long as its breadth. (The sjoecimen 
from which figs. 1 c and 1 e were drawn measures 51 mm. in 
length, and was captured on Stat. 146 ; figs. 1 a-\ h were drawn 
from the specimen secured on Stat. 159). 

But besides these five specimens of FetaUdhtm, I found still 
two smaller specimens of the same genus among the ' Challenger ' 
animals. One of these had been determined h.s 6'ergestes japonicus 
Bate, but is not referred to that species in his text. It was 
captured at Stat. 158, 7/iii., 1874, 1800 fathoms, thus near one 
of the above-named stations. It measures only 21 '5 mm. in 
length. Figs. 1_/ and 1 g show that its I'ostrum has the terminal 
process somewhat longer than in the large specimens ; the eyes 
are a little longer as compared with the length of their stalks, and 
these are proportionately somewhat broader, without a distinct 
knot at the end of the inner margin. There is no supra-ocular 
spine, but the hepatic spine is moderately developed ; the gastro- 
hepatic gi-oove is very distinct. No branchiae above trl.^ This 
small specimen belongs certainly to P. foUaceum Bate. 

Bate established (pp. 428-31) his Sergestes profundus on two 
specimens, both badly mutilated. He describes each specimen 
sepai'ately, beginning with one captured at Stat. 137, lat. 35° 59' S., 
long. 1° 34' E., dej)th 2550 fatlaoms. But, unfortunately, this 
specimen does not belong to Sergestes but to Petalidium ; the 
type is besides so mutilated that I should have preferx'ed to 
omit it, if it had not been described by Bate. It measures 
17"5 mm. in length. The rostrum is shown in fig. 2a; it is 
described by Bate : "It consists of a short fine point projecting 
horizontally for about one-fourth the length of the ophthalmopod, 
and is dorsally furnished on the crest with a small tooth." The 
eye-stalks are as in the sicpJl speci/nen from Stat. 158 just . 


described ; the posterior brancliiiB to a large degree are destroyed, 
and as to the other features, I refer to Bate's description (p. 429). 
I cannot say with certainty whether the animal is a very young 
specimen of P.foliaceum Bate, or belongs to an unknown form ; the 
rostrum deviates considerably from the type of P. folkic&um, but 
its shape presents a stage betv/een those in the adult and in a larva, 
nearly agreeing with that in a larval form mentioned below. 

In my earlier paper on Sergesies I described S. obesus Kr. 
{ = S. sanguineus Chun). I stated that it was a larva, a Masti- 
gopus, and added (p. 968) : " It is easily seen that this species 
cannot remain in the genus Sergestes, but whether it should be 
referred to Petalidium Ba,te, or a new genus should be established 
for its reception, is impossible to decide with certainty. The 
branchial plates recall the plates found in Petalidium, and there- 
fore I provisionally transfer it to that genus. ..." I had not 
seen any adult Petalidium, but suggested (p. 967) that the 
branchial plumes intei'preted by Bate as arthrobranchise are in 
reality pleurobranchise. I can now decide that S. obesus Kr. is 
the Mastigojms of Petalidium : the branchia3 pi-ove it ; besides, 
trl.^ in the Mastigopus is exceedingly elongate, with the basal 
part very thick, and in the adult form the basal joint of the same 
pair (fig. 1 e) is exceedingly thick, very much thicker than those 
of trl.^ and trl.^. Finally, the shape of the rostrum of S. obesus 
Kr, agi-ees nearly with that of the young Petalidiicm from 
Stat. 137 just mentioned. I described the branchicB of &'. obesus 
Kr., stating that in the largest specimen a rudimentary branchia 
was found above trl.* ; in the largest but one of the specimens I 
have now looked for this branchia, and found it. Finally, I stated 
that P. obesttm (Kr.) is "decidedly distinct irom P. foliaceiim Bate," 
and I maintain this opinion, founding it on the fact that P. foli- 
aceum does not possess any bi'anchia above trl.'' ; and it is very 
improbable that such a branchia begins to develop itself in the 
Mastigopus and disappears again in the adult. 

b. Sergestes H. M.-Edw. 

Serg. intermedius Bate, p. 383 (no figure). 

Bate mentions one specimen, 5 mm. in length, from the 
" China Sea, off Luzon." The specimen has been preserved ; 
according to the rostrum, the eye-stalks, and the ext. br. of urp., 
it belongs to *' Serg. brachyorrhos Kr.," which is the youngest 
Mastigopus of S. edivai-dsii Kr. (P. Z. S. 1896, pp. 963-64). 

Serg. prehensilis Bate, p. 385, pi. Ixxi. (Plate XI. figs. 4 a, 

Bate has examined one sj)ecimen from Stat. 236, lat. 34° 58' N., 
long. 139° 29' E., off Japan, 775 fathoms. The type, an adult 
male, has been preserved ; it differs from all other species known 
to me. It will be useful to redescribe its most essential characters, 
and two new figures are given (PI. XI. figs. 4 «, 4 6). The 

1903.] IX THE 'challenger' collectiox. 57 

ro.sti'um is rather long, directed forwards and considerably 
upwards ; its terininul portion is jiroduced into a spiuifonu 
piocess, and the upper margin of the rostrum has at the base of 
that process a sliarp angle as a rudiment of a spine ; the lower 
mai'gin of the rostrum between its base and the apical spine is 
sti'ongly convex, the upper margin nearly straight. Supra-ocular 
antl hepatic spines are wanting, the gasti'O-hepatie groove slightly 
developed. The eyes are lai'ge, considerably depressed ; seen from 
the side (fig. 4 a), they are somewhat longer than the whole stalk ; 
seen from above, their basal margin is very oblique (fig. 4 6), so 
that the interior margin of the distal joint of the stalk is as long 
as the outer margin of the eyes. The antenn. ped. with the outer 
margin of the first joint is a little longer than that of the two other 
joints together, and only a little shorter than their inner margin ; 
the second joint Avith the inner margin is three times longer than 
the breadth, and somewhat longer than the third joint, which is 
about two and a half times longer than broad. The antennal 
s({uama reaches nearly to the end of the antenn. ped., with its 
distal portion broad and the outer spine well developed. Mxp.^ is 
a little shorter than trl.^ The l'ranchi;e present a ti'ansition-form 
between those of b'. arciicus Kr. (PI. XII. fig. 1 c) and tS. robusivs 
Smith ; the pleurobianchial lamella above mxp.^ is very small. 
Of the bianchije above trl.^, br.^ is scarcely two-thirds as long as 
br. ; of those belonging to trl.^, br. is slightly longer than br\ 
and br.^ a little shorter than br} above trl.^ The ext. br. of vu-p. 
is almost five times longer than broad, and the haiiy portion of its 
outer margin is a little more than one- fourth of the total length. — 
Length 36"5 mm. 

This species occupies an intermediate position between S. arcticus 
Kr. and S. robitsius Smith. Bate's fig. 4, showing an antennide 
and an eye, is misleading, the antennular peduncle being too 
slender, with the basal joint too short, the third joint too long. 

Serg. japoxicus Bate, p. 387, pi. Ixx. figs. 1, 2. 

Bate enumerates three specimens from two localities : Stat. 232, 
lat. 35° 11' N., long. 139° 28' E., 345 fathoms; and Stat. 207, 
lat. 12° 21' N., long. 122° 15' E., 700 fathoms. All have been 
preserved, and belong to one species. In 1896 I wrote that 
S . j((j)onicus Bate must le identical with ^S'. mollis Smith (taken 
in the Atlantic, ofi' the United States), and gave reasons for 
my view. On comparing Bate's specimens with Smith's elaboi-ate 
description (Rep. U.S. Comm. Fish & Fisher, for 1882, p. 419, 
1884) and his figures (Rep. U.S. Comm. Fish etc. for 1885, pi. xx. 
figs. 3-5), I ai-rived at the same lesult. It should be especially 
mentioned that an examination of the bi'anchias showed the most 
complete agreement with Smith's description and drawing. For 
full information on S. japonicus Bate, I I'cfer, therefore, zoologists 
to the papers of Smith. 

As already mentioned, 1 found in the bottle with S. japonicus 
from Stat. 232 a tube containing a smaller animal determined 


to be S. jajyonicus and labelled Stat. 158. It is not mentioned 
in Bate's work. On a closer examination it turned out to be a 
specimen of Fetalidium, and it lias been described above. 

Serg. kroyeri Bate, p. 388, pi. Ixx. figs. 3, 4. (Plate XI. 
figs. 5 a, 5 b.) 

Bate established this species on one large specimen, from 
Stat. 170, lat. 29° 55' S., long. 178° 14' W., 520 fathoms. The 
specimen is very mutilated ; a new description with two figures 
(PI. XI. figs. 5 a, 5 b) is here given. 

The rostrum (fig. 5 a) is rather low, rounded above, with the 
upper front angle blunt and slightly projecting and the anterior 
niai'gin concave ; but it could not be settled whether the upper 
margin of the rostrum had been damaged or presented its natuial 
shape. Supra-ocular and hepatic spines are wanting, the gastro- 
hej)atic groove is strongly developed. The eyes (fig. 5 b) are 
large, somewhat depressed ; seen from above nearly as long as 
broad, slightly longer than the outer, and decidedly, but not 
much, longer than the inner margin of the stalk. On the upper 
side of the stalk, close behind the eye and near the inner margin, 
is seen a rather small, subcylindidcal, distally rounded process, 
directed obliquely forwards, inwards, and upwards ; it seems to 
terminate in an organ. In the antenn. ped. the basal joint is 
broad, with the outer margin a little shorter than that of the 
two following joints together, but only two-thirds as long as the 
inner margin of the same joints ; the second joint is stout, with 
the inner margin scai'cely three times longer than the breadth ; 
the third joint is stout, but yet considerably more slender than 
the second, scarcely three times longer than broad, somewhat 
longer than the outer and somewhat shorter than the inner 
margin of the second joint. The squama of the antennas seems 
to be nearly as broad at the distal end as in S. japonicus Bate, 
but it could not be seen whether the outer spine is developed. 
Mxp.^ and all trunk-legs are wanting. The pleurobranchial 
lamella above mxp.^ is small. The branchia3 above trl.^ have been 
broken ofi" on both sides ; of the branchiji3 above trl.*, br. is nearly 
three-fourths as long as br. above trl.^, while br.^ is slightly more 
than half as long as the same. The ext. br. of urp, with the 
apical part is wanting ; the branch seems to have been at most four 
times longer than broad, perhaps without marginal spine, and with 
the haired part of the margin unusually short. — Length 60 mm. 

The species is allied to t^. prehensilis Bate and S. robustics 
Smith, but it is easily distinguished from all species hitherto 
discovered by the process on the eye-stalks. 

Serg. atlaxticus H. Milne-Edw., Bate, p. 389, pis. Ixviii. &, 

In my earlier paper I wrote (p. 947) that " I am not convinced 
that all the specimens from the localities enumerated (p. 390) 
belong to S. atlanticus" and I produced some grounds for that 


opinion. The investigation of th© material presei'veil in the 
Britisli ]\Tu.seum pi'oved the cori'ectness of my disbelief, but, it 
must be admitted, to a degree not supposed. 

Of the specimens enumerated by Bate, the following have not 
been preserved in the British Museum : " Stat. 42, lat. 35° 58' N., 
long. 70° 35' W., 2425 fathoms," "length 25 mm."— and "On 
May 6-1 8th, 1876, in lat. 32° 41' N., long. 36° 6' W. . . one 
specimen ... at the surface ; and on the 7th of the same month, 
near the Azores .... two other specimens Avere taken at the 
sui'face." I can now state with cei-tainty that if the specimen 
from Stat. 42 lived near the bottom in that enoi'mous depth, it 
did not belong to S. atlanticus ; perhaps it was captured near the 
surface, but at all events the locality must be omitted as uncertain. 
IMost probably the specimens captured in May 1876 belonged to 
8. atlanticus. The specimens from the other stations enumei-ated 
by Bate belong to four species, and each station must be mentioned 

" Korth Atlantic .... Stations 62 and 63, on the passage from 
Bermuda to the Azores. Three specimens." In a bottle lal)elled 
"Between Bermudas and Azores" I found eight partly mutilated 
specimens of S. atlanticus. 

" Between Teneriffe and St. Thomas." In a bottle with the 
same locality, three specimens of S. atlanticus. 

"Station 320, lat. 37° 17' S., long. 53° 52' W., off Monte 

Video; depth 600 fathoms." Bate does not directly state the 
number of specimens, but he writes " Length 38 mm.," and the 
meaning is probably that he had one single specimen. In a bottle 
labelled " off Monte Video" I found six small and badly preserved 
specimens of >S'. atlanticus M.-Edw., and one large S2^ecimen of 
S. arcticus Kj*., and it is decidedly the last-named specimen which 
Avas procured from 600 fathoms. As to S. arcticus Kr., the 
student is referred to Kroyer's work, to the description and 
drawings in various papers of S. Smith, to notes in n}y earlier 
paper, and to some i-emarks below, in the description of iS. siviilis, 
n. sp., together with figs. 1 a-c on PI. XII. 

" Station 159, . . . south of Australia ; depth 2150 fathoms. . . . 
Three specimens. . . Length 43 mm." In the collection three 
large specimens are present, but they belong to 6'. arcticios Ki*., 
w'hich thus has been proved to be distributed through the deeper 
Atlantic, from the southeiii part of Greenland to lat. 38° S., and 
to south of Austi'alia. 

Bate enumerates still two deep-water stations, viz. Stat. 232, 
off Japan, and Stat. 173, off Matuku, Fiji Islands; he examined 
one specimen from each of these localities, but the animals belong 
to two new species to be described below. 

But before giving these descriptions I will sum up the results 
of the examination of the specimens referred by Bate to S. atlan- 
ticus H. Milne-Edw. ( =*S'.//-mi Kr.). The Copenhagen ]\[useum 
possesses some hundreds of adult specimens of S. atlanticus^ taken 
at a large number of places in the Atlantic, the Indian Ocean, 


and the western part of the Pacific, and all these were taken 
near the surface ; the ' Challenger ' specimens taken near the 
surface and referred by Bate to S. atlcmticus really belong to that 
species, but his specimens secured at the deep-sea stations 320, 
159, 232, and 173 belong to three other species. — I have found it 
unnecessary to attempt to point out what pai'ts of Bate's long 
description (and which of his drawings) can be applied to jS. atlan- 
ticus ; the zoologist is referred to the good desci'iption of Kroyer 
together with the additional notes in my earlier paper. 

Serg. similis, n. sp. (Plate XI. figs. 6 «-6 d.) 

The type of this species is the above-mentioned specimen on 
which Bate writes (p. 320) : "Stat. 232, . . . lat. 35° 11' N., long. 
139° 28' E., ofi" Japan ; depth 345 fathoms. . . . Length 50 mm." 
The species is closely related to S. arcticus Kr. The rostrum 
(fig. 6 b) is nearly oblong-triangular, a good deal longer than in S. 
arcticus (PI. XII. fig. 1 h) and directed more upwards ; the anterior 
margin of the scutum below the rostrum is strongly convex (fig. 6a) 
and protruding, while it is nearly vei'tical in S. arcticus (PI. XII. 
fig. 1 a). The supra-ocular and hepatic spines are well developed ; 
the gastro-hepatic gi'oove distinct. The eyes are nearly as in 
/S. arcticus, large, broader than deep, scarcely as long as broad 
(fig. 6 c), decidedly shorter than the outer margin of the distal 
joint of the stalk, and one-half shorter than its inner maxgin. 
The antenn. ped. (fig. 6 c) is about as in *S'. arcticus ; the outer 
margin of the basal joint is as long as the same margin of the two 
distal joints together ; the second joint is rather slender, its inner 
margin almost four times longer than its breadth and scarcely 
longer than the third joint, which is slender, about six times 
longer than broad. The squama of the antenna is moderately broad 
at the end, with the outer spine projecting beyond the terminal 
margin. Of the long appendages, mxp.^, trl.^, and trl.' are wanting, 
and the remaining thoracic legs are about as in S. arcticus. The 
branchise above trl.^ and trl.^ (fig. Qd) present excellent difierences 
from those in S. ai-cticus (PI. XII. fig. 1 c) ; br.^ above trl.^ is not 
quite half as long as br. above trl."^, and in spite of this not incon- 
siderable length it is semi-rudimentary, having only some posterior 
branches, nearly all short, and no anterior branches. Of the 
branchiee above trl.^, br. is long, four-fifths as long as br. above 
trl.'^ ; br.^ is more than half as long as br., very well developed, at 
least as long and more than twice as broad as br.^ above trl.' 
[In *S'. arcticus (PI. XII. fig. 1 c) — from a specimen obtained in the 
most northern part of the Atlantic — br.^ above trl.^ is about three- 
fourths as long as br. above trl.^, with well- developed branches 
on both sides ; of the branchiae above trl.'*, br. is only a little more 
than half as long as br. above trl.^, while br.^ is small, considerably 
narrower and shorter than br.^ above trl.''] The ext. br. of urp. 
has the apical portion broken off", but it is narrow, and seems to 
h% ve been a little more than five times longer than broad, thus 
sli ghtly narrower than in S. arcticus, but otherwise of the same 


shape aii'l with the marginal spine well developed. — Length 
54 mm. 

The species is easily distinguished from *S'. arctlcus Kr. hy the 
shape of the rostrum, together with the anterioi- margin of the 
scutum, and especially hy the branchiae above trl.^ and trl.' ; in 
all other features these two species aie closely allied. 

Serg. challengeri, n. sp. (Plate XII. figs. 2 a-2 n.) 

The type of this species is the specimen on which Bate writes 
(p. 390): "Station 173, Julv 24th, 1874; lat. 19^ 9' 35" S., 
long. 179= 41' 50" E. ; off Matuku, Fiji Islands; depth 315 
fathoms; bottom, coiul mud. One sjiecimen, male. Dredged. 
Length 24 mm." He referred it to *S'. atlanticus. I have the 
pleasure of appending the name of the renowned ship to this 
Sergestes, which is one of the most interesting species of Crustacea 
secured by the expedition. Unfortunately the single specimen is 
very mutilated. 

The rostrum (fig. 2 b) is luther low, short, obliquely triangular, 
turning somewhat upwards; its apex is iicute and very sliglitly 
produced. The supra-ocular spine is wanting ; the hepatic spine 
is rudimentary ; the gastro-hepatic groove (fig. 2 a) is deep, and 
the cervical groove very distinct. The eyes have been broken off, 
only the basal part of the stiUks being left. In the antenn. ped. 
(fig. 2 c) the outei- margin of the basal joint is a little longer than 
that of the two other joints together; the second joint is 
moderately robust, its inner margin a little more than three 
times longer than the breadth ; the third is slightly more than 
two and a half times longer than broad, a little shorter than the 
outer margin of the second joint, and only three-fifths as long as 
the inner margin of the last-named joint. The squama of the 
antenna is distally very broad (fig. 2 c), with the outer spine 
scarcely projecting beyond the terminal margin. Mxp.^ and trl.^- 
trl.' are wanting ; of trl.'^ the apical part has been lost, but these 
legs seem to have been a little longer than mxp.'-, and to be more 
slender than in S. arcticus, but otherwise not showing any 
difference of importance. Of the branchi;p (fig. 2 d), br. above 
trl.'^ and trl.^ are long and narrow ; &r.^ above trl." is as usual a 
lamella ; br.^ above trl.' is slightly more than one-third as long 
as br., especially with its anterior branches well developed ; of the 
branchi?e above trl.', ?)r. is alwut three-fourths as long a,s 6r. above 
trl.', while br.^ is proportionately large, nmch longer and broader 
than br.' above trl.\ and even more than half as long asb)-. above 
trl.' The ext. br. of urp. (fig. 2 1) has the apical portion 
wanting, but the branch seems to have been nearly five times 
longer than broad, with the marginal spine well developed and 
situated as in S. arcticus. — Length 23 mm. 

By the shape of the joints of the antenn. perl., the development 
of the branchiiP above trl.' and trl.\ and the shape of the ext. br. 
of urp., this species is related to ^S*. robustus Smith, .S'. preJiensilis 
Bate, and S. kruyeri Bate. But it is sharphj distiiKjuished from 


cdl other species hitherto known hy possessing an enormous multitude 
of luminous organs arranged regularly on the lower surface and 
near the lower lateral margins of the cephalothorax, on the six 
abdominal segments (figs. 2 of, i, h), on the sides of the shield 
(fig. 2 a), and on all the appendages preserved (figs. 2 c, d, e,/, g, h) 
with exception of the maxillulse and maxillse. The organs are 
easily seen ; they look almost similar to the eyes in Arane^e, and 
they differ much in size and direction. I have deemed it advisable 
to deal with the special arrangement and the structure of these 
organs in a separate section of this paper. 

Serg. dorsispinalis Bate, p. 394, pi. Ixxii. fig. 1. 

Bate does not state directly the number of specimens, but he 
had probably only one. The length is " 9 mm." and the locality 
"south of Australia, March 1874." An animal with a label of 
exactly the same contents was preserved in a microscopical pre- 
paration. It is a Mastigopus i-elated to S. arcticits Kr., &'. similis, 
n. sp., &c., but it could not be referred to any adult form. In 
the preparation the abdomen is seen from the side, and the 
cephalothorax essentially from below; the spine on the scutum 
on which Bate writes "just anterior to the [cervical] suture, in 
the median dorsal line, is a small, anteriorly directed tooth," is 
in reality the gastro-hepatic spine, which besides has been drawn 
in a position too remote from the front. Bate's figure is not 
correct in some other respects : in the antenn. ped. the second 
joint is about as long as the third, and both together somewhat 
shorter than the first ; the eyes reach to the end of the first joint ; 
the squama is longer than in the figure, reaching almost beyond 
the second joint of the antenn. ped. and distahy narrow ; the ext. 
br. of urp. is a little longer and conspicuously more narrow than 
in the figure, while its marginal tooth is indicated correctly. 

Berg, laterodentatus Bate, p. 395 (no figure). 

Bate has examined one specimen, measuring 8 mm. in length, 
and captured " south of Australia, March 1874." The type could 
not be found in the Museum. According to the description it is 
a Mastigopus belonging to the arcticus-grouTp, probably a younger 
stage of the same species as that to which the preceding larva, 
>S'. dorsispinalis Bate, belongs. 

Serg. nasidentatus Bate, p. 398, pi. Ixxii. fig. 2. 

Bate does not state directly the number of specimens, and 
presumably he had only one ; the length was 10 mm. and the 
locality the Pacific Ocean, between Valparaiso and Juan Fernandez. 
The type does not exist in the Museum. The species is a 
Mastigopus ; in my earlier paper I had already referred it to the 
same group as S. arcticios, and nothing further can now be added. 

Serg. diapontius Bate, p. 399, pi. Ixxii. fig. 3. 

Bate does not state the number of specimens, but presumably 


he had only one. The length is 18 mm. and the locality the 
Atlantic Ocean, AprO 7, 1876. In a preparation bearing the 
name of the species, and besides " Surface, 7 April, 1876 
Atlantic," one specimen is preserved : it must be Bate's type' 
but it measures only 16-5 mm. in length. In Bate's figure the 
armature on the dorsal line of the abdomen is not correct • on 
the third segment a spine has either been broken off or is rudi- 
mentary, the base being distinct ; the spine on the fourth segment 
is only half as long as that on the fifth, shorter than in the figure, 
and directed obliquely backwards ; on the sixth segment a%ery 
short sphie is visible. The basal part of the rostrum is somewhat 
ascending, the distal spiniform and horizontally porrected. The 
ext. br. of urp. has the outer margin hairy in about | of its 
length, and the marginal tooth is very small. The distal" part of 
mxp.' is very incorrectly drawn by Bate in his fig. 3 z ; it is four- 
jointed ; the third joint is short, much shorter^ than the fourth, 
and both together about as long as the second ; the first of these 
joints terminates below in two strong setiform spines, both some- 
what longer than the second joint, which terminates in two spines 
of the same length as the preceding jDair ; the fourth joint 
terminates also m two spines, somewhat, but not much, shorter 
than the four just mentioned. (I cannot understand how Bate 
drew his misleading figure ; it must prevent eveiy student of 
his Report from arriving at a correct judgment.) In the British 
Museum I have drawn tolerably accurate sketches of the rostrum 
the ext. br. of m^p. and the distal part of mxp.^ and I have 
compared them with a few specimens in the Copenhagen Museum 
previously determined and shortly described by me as the 2Iasti- 
gopus of S. penerinkii Bate, H. J. H. : I found the agreement to 
be so close, that I must consider S. diapontms Bate and S pene- 
rinkii Bate (the type of the latter form unfortunately is not present 
in the Museum) as two Mastigopiis-^\.&g%^ of the same species ; 
the type of S. penerinUi Bate measured only 8 mm. in leno-th 
and is a young Mastlgopus, while ^. diapontius Bate is the la?oe 
lai'va. ^ 

Bate describes S. diapontms on p. 399, S. loenerinkii on p. 418- 
and the name .S'. diapontius must therefore be accepted for the 
species. In my earlier paper I described the black-eyed adult 
form as S. penerinkii Bate, H. J. H., but I think that it must 
now be necessary to adopt the name S. diapontius, not only for 
the Mastigo^nis, but also for the mature form, which therefore in 
the future must be named ,S'. diapontius Bate, H. J. H. 

Serg. armatus Kroyer, Bate, p. 401, pi. Ixxiii. fig. 1. 

io-^?o^^T?'''''^''t*^'''^^ localities: one of them Ts " September 
12, 18/5, between Japan and Honolulu. South Pacific Ocean"- 
the second IS - Station 256, July 21, 1875; .... north of the 
Sandwich Islands ; depth 2950 fathoms." But in the collection 
I found a statement aberrant from both, viz. : " Surface. Japan 
to Honolulu, July 1875. Type." This bottle contained Iwo 


small specimens, but neither of them can be the type for Bate's 
figure. Both specimens belong to Group II. in my earlier j)aper, 
but neither of them belongs to S. armatus, both having on the 
rostrum a well-developed sub-basal dorsal spine, which is absent 
in S. armatus Ej". and in Bate's figure. I have been unable to 
refer the specimens, which measure about 8*5 and 11 mm., to any 
species known to me, and I thought it tiseless to describe and 
figure them. — One specimen from Port Jackson, Australia, the 
third locality in Bate's report, measures at most 6"2 mm. without 
the rostrum, which has been broken at the middle, but possesses 
a very fine svib-basal spine. It is so small and so badly preserved 
that a reference to any species has been impossible. 

Finally I found a specimen from "Sidney," determined by Bate 
as S. armatus, but not mentioned in his woi-k. It is only as long 
as the preceding specimen and impossible to determine. 

The result is that I have perhaps not seen the specimen figured 
by Bate, which may belong to aS*. armatus Kr., and, according to 
the explanation of the plate, measured about 10 mm. in length, 
while his specimens of this length examined by me disagree with 
his figure by possessing a sub-basal upper spine on the rostrum. 

Serg. edwardsii Kroyer, Bate, p. 403, pi. Ixxiii. fig. 2. 

Bate enumei-ates three localities. The first is " North Atlantic, 
April 1873 " : in the Museum I found a specimen labelled 
"14 April, 73, oS" Africa, surface," which most probably is that 
indicated in the text, and it belongs to 8. edioardsii Kr. The 
second locality is "Pacific Ocean, surface, September 1875": in 
the collection a specimen bearing the same inscription is>S'. edwardsii 
Kr. From the third of Bate's localities. Cape Yerde Islands, I 
found no specimen, but a specimen without locality and signed 
" type " is an adult specimen of S. edivardsii Kr. (That Bate's 
statement " Greenland [Kroyer) " is wrong here, and in almost all 
other places, has already been pointed out both by Ortmann and 
myself.) — Furthermore, I found two small specimens of S. ocidatus 
Kr., the Mastigopus of S. edioardsii Kr., which had been deter- 
mined by Bate as S. edioardsii and labelled " Aug. 23, 1873, 
lat. 2° 25' N., long. 20° 1' W., 100 fathoms," but these specimens 
are not mentioned in Bate's Report.- 

Bate's description of the characters of S. edioai'dsii Kr. is in- 
complete ; the reader is i-eferred to my earlier paper. 

Serg. rinkii Kroyer, Bate, p. 404, pi. Ixxiii. fig. 3. 

Bate mentions two localities : " New Hebrides, August 23, 
1874," and "South Pacific, 1875." From the first of these 
localities the anterior half of a specimen was present. Further- 
more, I found two specimens labelled " Oct. 19, 1875, S. Pacific, 
drawn," and one and a half specimen labelled "Oct. 18, 1875, 
surface" : both these localities are in all probability identical with 
the second one in Bate's Repoi't. It may be very possible that all 
these specimens belong to S. rinkii Kr., which is the Mastigopus 

1903.] IX THE 'challenger' collection. 65 

of S. arcticiis Kv., but liaving in Lniidon no matcrinl from the 
North Atlantic of S. rinkil Kr., for direct comparison of niinnte 
details, and some other species allied to S. arcticas being known 
from the Indian Ocean and the Pacific, I could not determine the 
larvaj enumerated with absolute certainty. 

One specimen from Cape York, detei-mined as &'. rinkii, but 
not mentioned in the Re^iort, is 6'. comic alum Kjr., in a stage a 
little younger than that figured by Bate as the last-named specias. 

One specimen, labelled " 25. 8. 73," determined as S. rinkii, 
but not mentioned in the Repoi't, measures nearly 7 mm. ; it 
belongs to S. ijenerinkii Jiate, the young larva of S. diaponilas 
Bate, H. J. H. 

Seeg. oculatus Kroyer, Bate, p. 406, pi. Ixxiv. fig. 1. 

Bate enumerates six localities. From Stat. 106 three specimens 
were obtained, from Stat. 257 one specimen, from Stat. 103 one 
specimen, finally from " September 12, 1875, South Pacific," one 
specimen : all six specimens correctly referred to the Ki'oyeriau 
species, which is the Masticjopiis of S. eclwardsii Kr. (compai-e my 
earlier paper). From the two remaining localities, viz. " North 
Pacific, near the Sandwich Islands, August 21, 1875," and 
"August 27, 1873 . . . ofi" St. Paul's Rock," no specimens could 
be discovered. 

Serg. ovatoculus Bate, p. 408, pi. Ixxiv. fig. 2. 

Bate gives the locality " The North Atlantic Ocean." In the 
collection I found three specimens with the label '"14 June, 1873," 
which agrees with my quotation from the text. These three 
specimens ai'e identical with *S'. avcylo'ps Kr., which is the 
Mastigojyihs of S. atlanticus IT. Milne-Edw. 

Serg. parvidens Bate, p. 409, pi. Ixxiv. fig. 3. 

Bate has the . following localities: " The tropical part of the 
Atlantic ; Pacific Ocean, north of the Sandwich Islands ; ofi" 
Sydney and Wellington, Australia." Just beloAv he writes: 
" Specimens of this species or vaiiety were taken during the 
passage from Teneriffe to St. Thomas" ; and in the collection one 
specimen from the last-named locality is present : it belongs to 
*S'. vigilax Stimpson, the Mastigojnts of S. vigilax Stimps., H. J. H. ; 
it agrees with Bate's description and figure, and it had already 
been pointed out by Ortmann and myself that *S'. 2>(i'''"^'idevs Bate 
belonged to *S'. vigilax Stimps. Examples from the other localities 
mentioned by Bate could not be found. — From " Cape York" two 
specimens determined as 6'. 2>(i^"videns were present : one of them 
is a young S'. ocidatus Kr. (the Mastigopus of «S'. edirardsvi Ki-.); 
the other is allied to 6'. incertus H. J. H., but is so badly preservett 
that a determination was impossible. In a preparation a small 
specimen from the " China Sea," determined as *,S'. parvidens, was 
preserved ; it seems to be S. ocidatas Kr., but it is in a very bad 

Proc. Zool. Soc— 1903. A^ol. I. No. V. 5 


Serg. corniculum Ki'oyer, Bate, p. 410, pi. Ixxv. fig. 1, 

Bate writes : " Cape York ; north of New Guinea ; ISTorth-west 
Pacific." In the collection I fovmd one specimen from Cape York 
labelled " type," furthermore, one and a half specimen from the 
same locality, all correctly I'eferred to the Kroyerian species. In 
a preparation I found a specimen determined as S. corniculuin 
from "ISr.W. Pacific"; it measures 10-4 mm. in length, but 
according to the shape of the ext. br. of urp., the eyes, &c., it is 
not that species but S. ancylops Kr., the Mastigopus of S. atlanticics. 
From the thii-d locality, " north of New Guinea," no specimen 
could be found. 

Serg. ancylops Kroyer, Bate, p. 413, pi. Ixxv. fig. 2. 

Bate has two localities : " New Hebrides ; Pacific, July 20, 
1875." I found one specimen from "New Hebrides" correctly 
referred to the Kroyerian form. — In three other tubes, specimens 
not mentioned in Bate's text were present ; they belonged to 
three other species, but are mutilated and could not be determined. 

Serg. longirostris Bate, p. 415, pi. Ixxv. fig. 3. 

Bate writes : " Mid Atlantic, April 1 876," and according to 
the following line he had one specimen. But in the collection I 
found three tubes, each containing one specimen, all determined 
as S. lojigirostris, and two of them from " N. Atlantic," while the 
third had no locality ; all three specimens are the young Mastigopits 
of S. corniculum Kr., H. J. H. The specimen without locality 
seems to be the type for Bate's figure of S. longirostris, but it 
measures scarcely 3 mm. in length, the rostrum not included, and 
in Bate's figure the eye-stalks are too long, the eyes too small, and 
the third joint of the antenn. ped. too short (it is in reality as 
long as the second). 

Serg. junceus Bate, p. 416, pi. Ixxvi. fig. 1. 

According to the text Bate has seen one small specimen from 
the "South Pacific Ocean." It could not be detected in the 
collection. But in my earlier paper I pointed out that it must 
be the young Mastigopus of S. tenuiremis Elr., H. J. H. 

Serg. longispinus Bate, p. 417, pi. Ixxvi. fig. 2. 

Bate has two localities: " Station 106 .. . Mid Atlantic" , . . 
and " Station 354 .... North Atlantic Ocean." The collection 
contained two specimens referred to this species and labelled 
" 23 Aug. 1873, 70 meters "; in the text we find "Station 106, 
August 25, 1873 ; lat. 1° 47' N., long. 24° 26' W. ; Mid Atlantic 
Ocean " • but in spite of the small difierence as to the date 
(probably originating from a missci'ipt) I am sui'e that one of 
the two specimens was captured at Stat. 106. The other 
specimen is probably from Stat. 354. Bate wi'ites : " the specimen 
from the ti'opical part of the Atlantic " and " that from the North 
Atlantic," which indicates that he had only one specimen from 


eacli of the two stations mentioned ; and I suppose, therefore, that 
both specimens have latei' on been put together in the sam« tube. 
Both specimens, which are in a very bad condition, belong correctly 
to S'. longispinus Bate (the direction of some of the dorsal spines 
on the abdomen is very characteristic), and this form is in reality 
the ^fast^gopus of S. conmtas Kr. (compai-e my earlier })aper, 
p. 953). 

Serg. pexerixkii Bate, p. 418, pi. Ixxvi. fig. 3. 

Bate records tlie length to be 8 nun. and the locality " North 
Atlantic Ocean." No specimen referred to this species could be 
found in the collection. But al)Ove I have mentioned a specimen 
of this species referi'ed by Bate to 6'. rinhii, and it may perhaps 
1)6 that desci-ibed by him as S. penerinklL He writes (p. 419) 
that the last-named " species bears a strong resemblance to 
/S'ergres^es r«iA;« Kroyer, but differs . . .". The specimen in question, 
referred by him to S. 7-inkil, measures nearly 7 mm. in leng-thand 
was captui'ed " 25. 8. 73," that is to sivy. Mid Atlantic. — In my 
earlier paper I have described the Masiigopits, and besides the 
mature form as *S'. peneruikii Bate, H. J. H. ; but above it is 
pointed out that this name must be cancelled as a synonym, and 
the species be named aS'. dlapontias Bate, H. J. H. 

Serg. fermerixkii Bate, p. 419, pi. IxxAd. fig. 4. 

Bate has examined one specimen, captured in the " Paciiic 
Ocean, lat. 24° S., long. 148° W.," and measuring 5 mm. The 
specimen is not to be found in the Museum collection. According 
to the figui'e it is a very young Mastigopus belonging either to 
»S'. diapontius Bate, H. J. H., or to a closely-allied species of the 
same group. 

Serg. loxgicollus Bate, p. 421, pi. Ixxvii. fig. 1. 

Bate enumerates two localities. The fii-st of them is " Soutli 
Atlantic Ocean, October 5, 1873 ; near Station 131 ; lat. 29° 35' 
S., long. 28° 9' W." In a preparation labelled ''5. 10. 73" I 
found the lai'ge specimen diuwn by Bate ; in a tube labelled 
"surface, at night, 5 Oct. 1873, South Atlantic," a small specimen 
was presei'ved. Thus both specimens are fi'om the fii'st locality 
in the text, and both belong to *S'. lo/igicoUas Bate, which is the 
Mastigopus of .S'. tenairemis Kriwei', H. J. H. In the large 
specimen di'iiwn by Bate the eyes present a thick yellowish Inyer 
around the black central part ; the dorsal spine on the sixth 
abdominal segment is exceedingly small. 

The second locality in Bate's text is " Station 295. November 5, 
1875 ; lat. 38° 7' S., long. 94° 4' W. ; South Pacific Ocean ; depth 
1500 fathoms; . . . Taken at night." In the collection ti specimen 
is preserved labelled " 5. Nov. 75, night, S. Pacific, surface." 
The determination is correct ; and the S2:)ecimen, which measures 
9"2 mm. in length, is evidently that mentioned by. Bat<? ; but 
the label proves that it was taken at the surface, wheiefore 



the depth recorded in the text is roost misleading. — In a third 
tube two specimens labelled 18/10 75 were preserved ; they had 
been correctly referred by Bate to his >S'. longicollus, but are not 
mentioned in his text. 

A preparation contained a specimen of the same species, 
labelled " Sergestes tenuiremis, South Atlantic, 6. 10. 73." It is 
not mentioned by Bate, who even writes (p. 421) that " No speci- 
men in the ' Challenger ' collection corresponds precisely with the 
description and figure given by Ki'oyer" of S. temoiremis. On 
the other hand, ^S*. tenuiremis Kr. and S. longicollus Bate are, as 
already stated, the same species. 

Serg. PRiECOLLUS Bate, p. 423, pi. Ixxvii. fig. 2. 

Bate has examined one specimen, measuring 25 mm., from the 
" IsTorth Pacific Ocean." The specimen could not be found. 
The shape of the eyes in Bate's figure shows that it has been a 
large Mastigo'jnijS. In my earlier paper I wi-ite (p. 958) : " is at 
least rather nearly related to S. corniculum Kr., from which it seems 
to difi"er by a somewhat different shape of the ext. br. of urp., 
and by having the fifth abdominal segment ' dorsally produced to 
a point.' " 

Serg. semiarmis Bate, p. 423, pi. Ixvii. fig. 1. 

Bate has two localities: "West Pacific Ocean "and "Station 
354, May 6, 1876 ; .... Mid North Atlantic." After the 
description he writes : " Observations. — A specimen (pi. Ixvii. 
fig. 2) very similar to the type was taken in the Atlantic . . . " ; 
and then he describes the specimen. I must suppose that he 
considered the specimen from the "West Pacific Ocean" to be 
the type of his S. semiarmis, and that the specimen described 
separately (p. 425) is that from ". . . May 6, 1876. . . ." Neither 
of the specimens could be detected in the Museum ; both are 
larvse, but a reference is impossible. In the collection I found 
one specimen determined as S. semiarmis and labelled "13 April 
1876, Atlantic, off" coast of Africa, surface"; it is a Mastigopus 
of S. diapontius Bate, H. J. H. 

Serg. l^eviventralis Bate, p. 425, pi. Ixvii. fig. 3. 

Bate had probably only one specimen, 7 mm. in length, from 
" North of New Guinea " ; it could not be found in the Museum. 
It is a young Mastigopus belonging to a species closely related to 
jS. arcticus Kr. 

Serg. spiniventralis Bate, p. 426, pi. Ixvii. figs. 5 & 6. 

Bate has only one locality : " North Pacific Ocean " ; the 
animal described measured only 3" 5 mm. in length, and some of 
its parts are shown in figs. 5 a, 5 Z, and 5 v, but fig. 5 v, repre- 
senting the ext. br. of urp., does not correspond at all with the 
description in the text (p. 427). The animal is not to be found 
in the collection ; it is a small Mastigopus related to S. vigilax 
Stimps. and allied species. Bate's fig. 6 a represents the head 


of " Sergestes sjjiniventrcdis var." from the " West Pacific " ; it 
has been suggested that the animal is allied to — or identical 
with — the Mastigopus of S. vigilax Stimps., H. J. H., but the 
specimen could not be found. The collection contained one 
specimen determined as ^S'. spiniventralis and labelled " Sidney to 
Wellington, 17. 6. 74"; it measures about 7'5 mm. in length, 
and the naked part of the outer margin of the ext. br. of urp. is 
slightly more than one- fourth of its length. The specimen is in 
all probability a young Alastigojyits of *S'. vigilax Stimps., H. J. H. 

Serg. trofuxdus Bate, p. 428 (no figure). (Plate XI. figs. 3 a, 

Bate has referred two specimens to this sjoecies. He describes 
each specimen separately: the first of them, from Stat, 137, 
belongs to Fetalidiiom, perhaps to P. foUaceum, and has been 
dealt with above. The other specimen, from " Stat. 300, 
December 17, 1875; lat. 33° 42' S., long. 78° 18' W. ; west of 
Valparaiso ; depth 1375 fathoms ; . . . . Ti-awled," is a real 
Sergestes, related to 6'. inous Faxon, but differing in the shape of 
the rostrum. Having removed the first-mentioned specimen 
from the genus Sergestes, I should think it justifiable to apply 
the name S. profunchis Bate to the last-named specimen, instead 
of proposing a new name. The animal is quite membranous, 
and is crimson everywhere — a fact proving that it lives in the 
depth of the sea, and that its colour has been dm-able to the 
highest and most unusual degree. The posterior part of the 
abdomen is Avanting, and the animal is on the whole so mutilated 
and flabby, that I would have omitted it if it had not been 
described by Bate ; but for that reason I have thought it necessary 
to add some notes with two figures (PL XI. figs. 3 a, 3 h). It 
agrees with ^S*. inous Fax. as to the membranous quality of the 
skin and the posterior branchite, but differs from it in the shape 
of the rostrum, which is of moderate length, with the upper 
mar-gin somewhat, and the lower margin paitly, sti-ongly convex, 
and distally it is pi'oduced in a moderately shoi't spine (PL XI. 
fig. 3 h). (Unfortunately I have not seen any specimen of the 
gigantic species S. inous Fax., and can therefore not decide whether 
the specimen described by Faxon had the rostrum uninjured.) 
Supra-ocular and hepatic spines are wanting. The eyes (PL XI. 
fig. 3 a) are black, compaiutively small, somewhat shoi'ter than 
broad, not half as long as the eye-stalks, and not broader than the 
distal end of the stalks. In the antenn. ped. the outei' margin 
of the first joint is almost as long as that of the two distal joints 
together, therefoi-e somewhat shorter than their inner mai'gin ; 
the thix'd joint is a little shorter than the inner margin of the 
second, and seems to be about thi'ee and a half times longer than 
deep. The squama does not reach the end of tlie antenn. ped., 
and the outer distal spine is well developed. Accoi'ding to Bate 
the part preserved measures 18 mm., and the probable length of 
the entire animal is 24 mm. 


Serg. ventridentatus Bate, p. 431 (no figure). 

Bate gives the locality " north of the Sandwich Islands " and 
the length " 7 mm." A specimen labelled in accordance with the 
text is preserved in balsam ; it is a young S. oculatus Kr., the 
Mastigopus of S. edwardsii Kr. 

Serg. utrinquedens Bate, p. 433 (no figure). 

Bate gives the locality " North Pacific Ocean," and the length 
" 3"5 mm." No specimen could be found in the collection. Bate's 
specimen is a very young Mastiyopits ; in my earlier paper I 
have placed it near S. comiculum Kr., but a final interpretation 
is impossible to me. 

Serg. DissmiLis Bate, p. 437 (no figure). 

In the collection one specimen, from Cape Verde Islands, is 
preserved ; it is certainly the type described by Bate. In my 
earlier paper I had determined it as one of the larval stages of 
aS'. arctic'us Kr., but this is not correct. The rostrum is slightly 
more than one-third the length of the eye-stalks, its basal part 
somewhat ascending, with a very small spine on the upper angle, 
its distal part is a slender horizontal spine. The hepatic spine is 
short. The eyes are only a little higher than the distal end of 
the eye-stalks, and these increase gradually in thickness from the 
base outwards. In the antenn. ped. the distal half of the first 
joint has the margins nearl}^ parallel, and the two other joints ai-e 
a little thicker than in the cori-esponding Mastigopus of S, arcticus. 
Fourth to sixth abdominal segments each with a very small dorsal 
spine directed backwards ; the spine on the sixth segment is the 
longest. The ext. br. of urp. is almost, but not quite, five times 
longer than broad. All these characters agree with those found 
in a Mastigojjus of ^S'. mediterraiieus H. J. H. preserved in the 
Copenhagen Museum ; and sketches drawn in London of the shape 
of the distal part of the squama and the telson agi^ee also with 
the last-named form. I must therefore consider S. dissimilis Bate 
as identical with *S'. mediterraneus H. J. H. The result is that 
the last name must be withdrawn as a synonym, and the species, 
of which the sut-adult stage has been described in my earlier 
paper, mvist be called S. dissimilis Bate, H. J. H. 

II. Some Residts of the Investigation. 

In my earlier paper on Sergestes I have paid as much attention 
as possible to the animals described by Bate ; in nearly all cases 
I was able to state whether the specimen was an adult form or a 
Mastigoptis, and several of the species were interpreted. After 
the study of the material preserved in the British Museum, I have 
now been able to confirm most of my earlier statements, to 
interpret an additional number of the specimens mentioned by 
Bate, and to correct two faults committed by myself. I had 
ei'foneously referred S. dissimilis Bate to S.^ arcticus Kr., instead 
of identifying it with S. mediterraneus H. J, H. (see above). 

1903.] IX TUE 'challexger' collectiox. 71 

Furthermore, I had divided a number of species, enumerate! on 
p. 949 as Group I. A. h. fy, into two sections, according to 
difierence in the thickness of the distal joints of the antenn. ped.; 
but a study of the types of S. 2>^'6hensilis Bate and S. kroyeri 
Bate showed that Bate's drawings of the antennul;ie of these 
species ai-e incoirect and misleading, wherefore my ari'angement 
of them wtvs wi-ong. 

It may be useful to put together the alterations and additions 
which may now be accepted in the Consj^ectus on pp. 949-51 in my 
eai'lier paper. In Group I. the following particulais must be 
added or altered : — To *S'. atlanticus H. M.-Edw. belongs only a 
part of S. atlanticus sens. Bate, besides the foi-m i-efeixed by Bate 
to S. ancylops Kr. From S. arcticus Kr., aS'. dissimilis Bate must 
be removed, and the latter species is to be established separately 
with 6'. mediterraneus H. J. H. as a synonym ; f ui-thermore, some 
of the specimens referred by Bate to S. atlanticus belong to 
S. arcticus. Near S. arcticus Kr., must be insei-ted S. similis 
H. J. H., established on one of Bate's specimens of S. atlanticus. 
S. prehensilis Bate and S'. kroyeri Bate must be removed fi-om 
their place and inserted above near S. japonicus Bate, together 
with aS. 2^rofuvdus Bate, in its new restriction, and S. challengeri 
H. J. H., established on one of the specimens referred by Bate to 
S. atlanticus. S. longirostris Bate must be withdi'awn as being 
a Mastigopus to S. cornicidum EJr., H. J. H., and S. corniculum 
sens. Bate is the same species. As uncertain remain : S. dorsi- 
spinalis Bate, ^S*. laterodentatus Bate, S. nasidentatus Bate, S. Icevi- 
ventralis Bate, *S'. rinkii Bate, ? KJr., all larval forms belonging to 
species related to S. arcticus Kj.\, or perhaps partly belonging 
to S. arcticus itself ; furthermore, the larvas S. prmcollus Bate, 
S. utrinquedens Bate — both at least rather closely related to 
S. cormculu/m Kr., H. J. H., — and S. semiarmis Bate. 

In Group II. there is hardly anything to alter, but some additions 
to make. To S. edwardsii Kr. belongs *S'. edwardsii Kr., Bate, 
S. oculatus Kr., Bate, S. intermedius Bate, and S. ventridentatus 
Bate. S. jjenerinkii Bate must be cancelled as synonymous with 
aS'. diapontius Bate ; and the adult form described as »S'. penerinkii 
in my eai-lier paper must be named ;S'. diapontius Bate, H. J. H. 
The rest of Group II. remains unaltered. S.fermerinkii Bate, 
S. spiniventralis Bate, and the species referred by Bate to 
S. armatus Kr. coidd not be interpi'eted. 

Besides, the investigation has yielded some results on the 
bathymetrical and geographical distribution of some of the species. 
It has been proved that the large specimens (exceeding 30 mm. 
in length) referred by Bate to S. atlanticus Kr, are deep-sea 
forms belonging to other species : >S'. atlanticus is common near 
the surface; according to Ortmann it has been taken in the 
intermediate net from 700-500 m., but it does not live in the 
greater depths of the sea. S. arcticics Kr. has a very wide 
geographical range, through the deeper to very deep tracts of the 
Ocean (see above); S. japonicV'S Bate has been captured in 
the northern part of the Atlantic and the northern pai-t of the 


Pacific. ^S'. atlanticus H. Milne-Edw., S. edwardsii Kr., S. vigilax 
Stimps., H. J. H., S. tenuiremis Kr., H. J. H., and .S'. corniculum 
Kr., H. J. H., have been proved to be distributed through the 
tropical and subtropical parts of the Oceans almost around 
the globe, viz., from lat. 23° N., lat. 32° N., or even lat. 42° N. 
in the Atlantic, throughout the Indian Ocean to Australia, New 
Hebrides, and " South Pacific." 

III. Limiinous Organs in Sergestes challengeri, n. sp. 
(Plate XII.) 

The luminous organs briefly mentioned above are generally 
easy to discover ; each resembles a very convex, vitreous, faintly 
yellowish lens, which is circular and sharply defined. They differ 
much in size, some of them being very small and many pro- 
portionately large. It may be advisable, first, to give a con- 
spectvis of the oi'gans observed on the single and tmfoi'tunately 
very mutilated specimen, next to add some remarks on their 
distribution, then to describe their structui-e, finally to compare 
them with luminous organs in other Crustacea. 

Conspectus of the Organs observed. 


On each side of the scutum a row with four organs. 8 

On the clypeus one oi'gan 1 

On the lower side of the head one unpaired organ 

and one near the lateral mai'gin 3 

On the lower surface of the thorax and on its inf ero- 

latei'al anargins 26 

On the lower side of the third joint of the antenn. 

ped. one organ 2 

On the lower side of each squama four oi-gans 8 

On the mandible and its palpus two organs 4 

On the first maxilliped two organs 4 

Oil the second maxilliped five organs 10 

On the first trunk-leg three organs 6 

On the lower surface and on the lateral wall of the 

first abdominal segment , 6 

Do., do. of the second abdominal segment 6 

Do., do. of the third abdominal segment 4 

Do., do. of the fourth abdominal segment 3 

On the latei'al wall of the fifth abdominal segment. 2 

Along the median line of the sixth abdominal segment. 6 
On the outer side of the basal joint of each of the 

pleopods one organ 10 

On the peduncle and on the inner branch of each 

uropod two organs 4 

On the lower side of the outer branch of each ixropod 

two organs 4 

Total number of organs ... 117 


The eye-stalks, the moxillipeds, and the four posterioi' pairs of 
the trunk-legs have been broken ofi". I am convinced that at 
least most, and perhaps all, these appendages possess some oi-gans, 
and the lowest number the species possesses must therefore be 
about 150 ! 

On the distribution and direction of these organs the following 
remarks may be offered. The four orgjins on each side of the 
scutum are arranged i-ather close in a longitudinal i"ow situated 
on the ridge bordering the branchial cavity above (fig. 2 a). The 
organ on the clypeus is lai-ge : one oi'gan is situated on the 
segment bearing the antennulre, and one on the lower surface of 
the head near the lateral margin a little in front of the mandible ; 
these four oi'gans look essentially dow^lwaI■ds. The arrangement 
of the organs on the three posterior thoracic segments can be seen 
in figs. 2 i and 2 d. It is observed that two large organs are 
placed at the lower margin of br. above trl.^ and of br.^ above 
trl.'' These organs look outwards and downwards, ard the part 
containing the glandular mass etc. behind the lens is somewhat 
protruding, which produces an aspect as if these organs had been 
inserted on the end of a kind of rather thick, short stalk. The 
remaining oi-gans on the segments mentioned are found on the 
lower surface (fig. 2 i) ; those placed at the inner angle of the 
legs are small oi- very small, while some in the median line are 
large. In fig. 2 i fifteen thoracic oi'gans have been drawn ; the 
remaining eleven thoracic organs ai-e situated on the antei-ior 
segments and arranged in a rather similar way. The four organs 
on the lower surface of each squama of the antenna are arranged 
for some distance in a row ; two of them are seen through the 
sfjuama in fig. 2 c. The mandible (fig. 2 e) has one organ below 
at the antero-inferior margin near the insei'tion of the palp ; 
another oi'gan is seen on the lower surface of the first joint of the 
palp near its distal end. The first maxilliped (fig. 2/) has on 
the upper side one organ just at the origin of the exopod, and one 
on the lower side of the following joint of the endopod ; the fii'st- 
nained organ looks forwards and a little inwards (fig. 2 g), and 
the upper mai-gin above it is produced nearly as a lamella, over- 
lapping the major part of the lens wdien seen from above (figs. 2 (j 
<fc 2 n). Of the oi-gans on the outei' — in the natural position of 
the appendage in reality the lower — surface of mxp." (fig. 2 k), 
tha.t at the base of the third joint is very small and the others 
large ; the oi-gan on the first joint looks essentially downwai'ds, 
and is " stalked," as the above-described organs near the lower 
margin of the posterior branchiae. Of the three organs on trl.\ 
two are placed on the inner side of the long fourth joint, one near 
the base and the other near the distal end ; the third organ is 
situated on the anterior side near its end. Each of the five 
anterior abdominal segments has a lai'ge organ on the lower pai-t 
of the antei'ior margin of the lateral plate (figs. 2d & 2 1) ; it 
looks forwards anrl somewhat downwai-ds, besides sometimes a 
little outwards. Each of the two antei'ioi- segments has besides 


on the lower surface (fig. 2 i) four organs, one of which is large 
and two of the others very small ; on the third and fourth 
segments these organs are gradually reduced in number, and none 
of them are found on the fifth. The sixth abdominal segment 
has a median row of six organs, which are seen in fig. 2 k with 
the exception of the first, this being hidden by the lateral wings 
of the preceding segment. The very short basal joint of the 
peduncle of each of the pleopods has on the outer side a small 
organ, looking outwards and at least sometimes a little down- 
wards. Each lu'opod has one organ on the pedvmcle, situated on 
its inner surface near the lower margin and close to the base, 
besides one organ on the interior (lower) surface of the inner 
branch near its base ; finally, two organs on the interior (lower) 
surface of the outer branch, one of them near the middle and the 
other on the distal nari'owing part. 

From the preceding description it is seen that most of the organs 
look downwards, a smaller number somewhat outwards or forwards, 
rather few almost totally outwards, and none upwaixls, with 
the exception of one on the first maxilliped. With the exception 
of the few lateral organs on the scutum, all the others are found 
on or near the lower surface of the body and on the appendages. 

The structure of the organs is very intei'esting, and very 
difierent from all hitherto known in any invertebrate animal. I 
have examined more closely three organs, viz., that situated on 
the third joint of the first maxilliped, one of the "stalked" 
organs near br. above trl.^, and one from the antero- inferior 
margin of the fourth abdominal segment. The two last-named 
organs have been cut off, most of the adhering tissue removed 
by dissection, and the organs examined with moderately high 
magnifying power. I ha,ve found no difi"erence of any importance 
between the three organs from such distant parts of the animal's 

The organ taken as type is that from the infero-lateral margin 
of the thorax ; it has been drawn (fig. 2 ni) in optical vei'tical 
section. The skin forms a chitinous, large, and very thick biconvex 
lens (a), which is vitreous and a little yellowish ; the major 
central part is covered by a rather thin limpid layer («^), but this 
layer I could not perceive on the two other organs examined. 
The lens is circular in outline ; its diameter is about two-thirds 
as long as that of the inner portion of the organ at its thickest 
part. The inner side of the biconvex lens is covered by a large 
and rather thick concavo-convex lens (6), which is somewhat 
thinner than the outer lens, but with the diameter a little longer 
than that of the same ; the lateral margin of the inner lens is 
oblique, very broad, touching the external chitin around the 
outer lens. This inner lens, which consists of two layers, is 
homogeneous, vitreous, and slightly greyish ; but the difference 
between the colour of the outer and the inner lens is in the figure 
purposely a little more strongly marked than in nature. These 
two lenses remind one of optical instruments in which the lens is 


composed of crown-glass and flint-glass. Behind the inner lens 
is found a thick layer of glandular cells {d), A\hich are light 
greyish, very lai'ge, and most of them elongate, I'adiating towards 
the centre of the outer lens. The diameter of this layer is some- 
what larger tlian that of the inner lens ; and when the luminous 
organs are examined in their natural position with a strong 
pocket-lens, this layer can often be seen through the skin as a 
whitish ring around the lens. Between the layer of large cells 
and the inner lens a thin layei- (c) seems to exist, but its quality 
could not be made out, and I do not venture to pi'ojoose any 
hypothetical explanation. Behind the glandular layer is seen 
another (e), Avhich is yellowish, with numei-ous ti-ansverse fine 
stiipes, and without ti-ixce of cellular sti-ucture ; it is i-ather thick 
in the middle, but thin around the sides of the glandiiiar layer. 
The internal surface of the organ seems to be covei'ed l)y a thin 
layer (/') of connectiv^e tissue. That the posterior layer with the 
ti-ansvei'se stiipes is — as in the EuphaiisiicUe — a I'eflector can be 
taken for granted. But it is impossible to decide wdiether the 
light is pioduced by the glandular layer or by the inner lens. 
AVhether the thin layer enveloping the whole organ is pigmented 
or not cannot be seen on this old material, which has been 
pi'esei'ved twenty-eight yeai-s in spiiit ; that the oi-gans ai'o 
immovable scarcely needs to be stated. Future investigation of 
living animals and of sections of fresh inateiial must elucidate 
whether the organs are especially innervated or not, and, besides, 
fill up the other gaps in the interpi'etation of the function and 
structui-e of the layers in these compound structures. 

A bi-ief compai-ison of the luminous organs in Sergestes 
challengeri with those in other Crustacea is not without interest. 
Of animals belonging to that class, luminosity has been observed 
in some Copepods, a few Ostra.cods, Euphausiidpe, and one 
macrurous Decapod. Giesbi'echt has publislied a thorough and 
interesting study : " Ueber das Leuchten der pelagischen 
Copepoden und das thierische Leuchten im Allgemeinen" (Mitth. 
zool. Stat. Neapel, 11 B., 1895, pp. 648-689). "He has examined 
a small nuinlier of jielagic Copepoda — necessaiily I'estrieting 
himself to all the luminous forms which he could jjrocure in a 
living state in the Gulf of Ka.ples — showing that these animals 
possess a number of small dermal glands, the secretion of which 
produces the luminosity when it, by exhaustion from the glands, 
comes in contact with the suirounding water. In a few Ostracoda 
a brilliant luminosity is produced in a similar way from glands in 
the labrum ; it Avas already suggested by G. W. Miiller in 1890, 
and has since been observed and published by another author. 
Furthermore, I can mention that during a voyage in the Indian 
Ocean, Dr. Th. Mortensen met with a vast number of a pelagic 
Ostracod which showed a most brilliant light, and he obserAefl 
how this was produced. Finally, the present writer has obser\-ed 
nejirly the same in a number of a lai-ge Ostracod which had 
been procured in Davis Strait. Lute in the evening I observed 


luminoiis points in recently sieved bottom material from 
318 fathoms, and undertook instantly some manipulations with 
the animals : the luminous fluid came from the head, probably 
from the large labrum, and flowed backwards between the shells, 
illuminating brilliantly the space between them, especially on the 
ventral side. As to the Entomostraca, we can therefore not speak 
of " luminous organs " in the common sense of the word, the light 
being always produced in the above- described way. 

In almost all Euphausiidee real and highly- developed luminous 
organs are found, but they differ in structure very much from 
those in Serg. challengeri. The reader is referred to Sars's 
Report on the ' Challenger ' Schizopoda, and to Chun, " Leucht- 
organe und Facettenaugen " (Bibliotheca Zoologica, Heft xix. 
Lief. 4, 1896). The highest number of organs met with in this 
family is only ten : one on each eye-stalk, two on each side of the 
thorax near its inferior margin, and one in the median line of 
each of the four anterior abdominal segments. The light is 
produced by the " Streifenkorper " (Chun) — "a flabell if orm bunch 
of exceedingly delicate fibres, exhibiting in fresh specimens a most 
beautiful iridescent lustre" (Sars, op. cit. p. 71) — situated a little 
behind the centre of the organ in a mass of large cells ; a biconvex 
lens, which is present in the organs with the exception of those on 
the eye-stalks, is internal, while the outer chitinous skin is thin ; 
a reflector is developed nearly as in Sergestes. 

The Danish zoologist Cand. mag. Ad. S. Jensen has directed 
my attention to a book by a French author, and lent me a 
German translation : ' Die Leuchtenden Tiere und Pflanzen. Yon 
Henri Gadeau de Kerville. Aus dem Franzosischen iibersetzt 
von W. Marshall, 1893.' In this popular treatise I saw that the 
' Talisman ' had captured a deep-sea shrimp with numerous 
luminous organs. I attempted in the ' Zoological Record ' and 
elsewhere to discover where that animal had been described, but 
in vain, and I will therefore reprint the passage in question from 
the German book : — " Wahrend der wissenschaftlichen Expedi- 
tionen des Talisman fing man in einer Tiefe von 500 m. einen 
langschwanzigen Krebs {Acanthephyra pellucida A. Milne - 
Edwards), welcher ein lebhaftes Licht um sich zu verbreiten im 
stande war und zwar mittels folgender verschieclener Leucht- 
organe; erstens befand sich eins am Yorderrand einer Deckschuppe 
der Augen, zweitens verlief eine lange leuchtende Linie am 
Aussenrand des Tarsus des fiinften Beinpaares, an dessen innern 
Basis sowie an der des vorhergehend en Beingiiedes sich weiter 
leuchtende Flecke befanden, drittens lag^n ganz ahnliche Flecke 
an der Basis des zweiten Gliedes des dritten und vierten Bein- 
paares und ebenso je einer an der Basis des Tarsus derselben 
Gliedmassen, viertens sah man einen langen Fleck an der Basis am 
Endgliede des hintersten Kieferfusspaares, fiinftens verlief ein 
schimmerndes Querband iiber die Hiifte des hintersten Thorakal- 
f usspaares, sechstens war eine Doppelreihe glanzender Punkte an 
jedem Gliede der Aussengeissel der Brustfiisse, sowie an dem. 
ausseren Blatt der BaucMiisse vorhanden, siebentens zeigte sich 


eiiie (loppelte L'nie leuehtender Puiikte ent.laug der aiisseren 
Geissel der luneufiihler nml achtens verlief eine iin hiuteren 
Telle zusammeuhaiigende, iiu vordei'eii in Puukte aufgelciste Linie 
parallel zum Uutenuude des Riickeupanzei's." The arrangeineut 
of tlie organs in Acanthephyra pelliocida is rather different from 
that in Sergestes chcdlengerl, but it shows yet more resemblance 
to it than to that in the Euphausiidie. Both in *b'. challengeri 
and in A. pellucida an astonishingly high number of organs exist, 
but as to the structure of the organs in the last-named form 
unfortunately nothing is known. I suppose that they are renl 
compound organs, not dermal glands aa in the Entomostraca. 

If we look for comparison between all luminous animals, it will 
1)6 found that only some deep-sea or pelagic fishes and two 
Cephalopoda of the genus Jlistloleathis (according to Verany and 
Joubin) possess a number of ryal organs which can be compared 
with that found in iSe7'g. challengeri and Acanth. pellitclda. 

I have looked for luminous oi'gans in all adult species of 
Sergestes known to me and in Acaii.ihephyra pLirpn,rea A. M.-Edw., 
but the result was absolutely negative. It is a very curious fact 
that about 150 very compound organs are found in one sj^ecies of 
Sergestes, while they seem to be quite absent in all other species 
hitherto known of the geniis. It may be added that the luminous 
species does not deviate from some of the other foi-ms in any other 
character of importance : it belongs, even within the genus, to a 
group which contains several deep-sea. forms closely allied to e;ich 
other. Considered in this light, the existence in one species of 
about 150 compound organs seems to me a most astonishing 

Su2)p>lementary Note. 

In the preceding section it has been mentioned that among tlie 
Cephalopoda two species of HistioteiUlus possess a large number 
of compound luminous organs. My frieiuls Prof. G. B. Howes 
and Rev. Th. R. R. Stebbing have very kindly dii'ccted my 
attention to two papers, which contain some additional knowledge 
of the same topic and may be quoted here. W. E. Hoyle (Mem. 
& Proc. of the Manchester Liter. & Philos. Soc. vol. xlvi. part vi. 
1902) points out and describes the structure of twenty-nine 
luminous organs in Pterygioteidhis margaritifera, but this numbei", 
though rather high, is yet considei'ably lower than that in 
Sergestes challengeri. C. Chun (' Aus den Tiefen des Weltmeeres,' 
Jena, 1900) describes and figures (p. 532) the arrangement and 
the colours of twenty-foui- luminous organs in '■^ Enoploteuthis 
diadema Ch., n. sp." Furthei'more, he writes (pp. 532-33) : 
"Ahnliche, wenn audi etwas kleinere Organe, besetzen bei 
Vei'ti'etern der Gattung Calllteathis <lie gauze Kiirperobei'llache 
von den Armen bis zu den Scliwanziiossen. Die Bauchseite ist 
auch liier wieder reichliclier mit ihnen a.usgestattet, als die 
Riickenfiiiche." An acconqjanying figui'e of a '' CalUteuthis n. sp.," 
seen from the ventral side, shows a nuinl:er of organs consideraljly 
surpassing that in Serg. challengeri. 


Plate XL 

1. PetalicUum foHaceum Bate. p. 34. 

Fig. 1 a. Rostrum of a specimen from Stat. 159, from the side. 

1 b. Eye and left antennular peduncle of the same specimen, from above. 

1 c. Rostrum of a specimai f/om Stat. 116, from the side; X 5. 

1 d. Eyes of the same specimen, from above ; slightlj' more than X 5. 

1 e. Basal part of mxp.^ — mxp."* and trl.i — trl.'', together with the branchial 
apparatus of the last-named specimen ; X 5. br., rudimentary pleuro- 
hranchia to mxp.^; ep., epipod on raxp.2 with its branchia j I., four 
pleurobranchial lamella? belonging to mxp.-* and trl.' — trl.^ 

ly. Lateral view of the front part of the scutum and the eye of a young speci- 
men — measuring 21'o mm. in length — from Stat. 158. 

Iff. Rostrum and eyes of the same specimen, from above. 

2. Fetalidmm sp. p. 55. 

Fig. 2 a. Front portion of the scutum of a young specimen from Stat. 137, described 
by Bate as Sergestes profundus Bate. 

3. Sergestes profundus Bate. p. 69. 

Fig. 3 a. Front part of the scutum, eye, and antennular peduncle of the specimen from 
Stat. 300; X15/2. 
3&. Rostrum of the same specimen. 

4. Sevffestes preJiensilis Hate. p. 56. 

Fig. 4(7.. Rostrum, eye, and base of the antennular peduncle, from the side. 

4 b. Front end of the scutum, ejes, jpeduncles of the antennulie and squama?, 

from above. 

5. Sergestes Icroyeri Bate. p. 58. 

Pig. 5 a. Rostrum, from the side. 

5 b. Front end of the scutum, left eye, and the peduncles of the antennulaj, from 

above; X 4. 

6. Sergestes similis, n. sp. p. 60. 

Fig. 6 a. Front end of the scutum, left eye, and peduncle of the antennula, from the 
side ; X 14/3. 

6 b. Rostrum and supra-ocular spine, from the side; X 12. 

6 c. Front end of the scutum, eye, peduncles of the antennulte and squamaj, 

from above ; X 5. 
Qd. Branchia; above trl.2— trl.-i ; X 11/2. 2br., first branchia to trl.2; SI., 

branchial lamella to tr]."; Sbr., first branchia to trl.'^; 3 br.^, second 

branchia to trl.'* ; &c. 

Plate XII. 

1. Sergestes arcticas Kr. p. 60. 

Fig. 1 a. Front end of the scutum, eye, and basal portion of the antennular peduncle, 
from the side ; X 9, 2. 
1 b. Rostrum and supra-ocular spine of the same specimen ; X 11. 

1 c. Brauchiffi above trl.^ — trl.-i ; X 8. The lettering as in fig. 6 d on the pre- 

ceding Plate. 

2. Sergestes challengeri, n. sp. p. 61. 

In the following figures o signifies luminous organs. 

Fig. 2 a. Scutum, with the basal parts of the eye-stalk, antennula and antenna ; 
X 5. 

2 b. Front end of the scutum, basal part of the eye-stalk, and antennular 

peduncle; X 17,2. 
2 c. Front end of the scutum with the anterior appendages, from above ; X 9. 
Of the eye-stalks only the basal ioint remain.s ; of the left squama the 
distal half is omitted; on the right squama the two distal ones of the four 
luminous organs are seen through the plate. 


Tig. 2d. Tlii>lias;il ])iirt-< of tlu' tliR'O i)i)stcvi<ir thoracic Icfrs, tlic hvaiicliiipaliovo trl.- — 
trl.', ami the lower i)art ot' the two iiiitcrior abiloiiiinal sijiniciits, Iroin the 
siilc; X 11. The lettering of the braiichiic a.s in tig. Od on the nveceilin"- 

2 e. Left niandibli', from helow ; X 13. 

2/. Left tirst niaxilliped, from helow ; X 13. 

2 (/. Middle part of the tirst maxillijied, from ahove ; X it. o', iipjier provimal 
himiiK^iis orjran ; o-, distal luminous organ seen througli the tindt)pod. 

2Ji. Left second maxilliped with its ejiijiod and hranchia, from the outer side ; 
X 13. 

2 i. The three posterior thoracic segments and tlio two anterior ahdominal 
segments, from helow, showing 31 luminous organs; X !>. Of the 
l)h'opods, only the basal joint — with its lumiiu)us organ — lias heen drawn. 

2 k. Sixth ahdominal segment with the hasal ])art of the telson and the uro])()ds, 
and the posterior part of the fifth abihmiinal segment with a part of the 
pleoi)od ; X 13/2. 

2 1. Exterior branch of the left uropod, from the outer side; X 17,2. The 
apical part is wanting. 

2 m. Luminous organ from the side of the thorax near br. ahove trl.^, seen in 
oi)tical vertical section; X 180. a, chitinous lens; a', its thinner 
external layer; b, inner lens; c, thin layer between the inner lens and the 
thick glandnlar layer d; c, reflector; /", enveloiMiig thin layer. 

2«. Sketch of till' luminous organ from the ujiper side of the tirst maxilliped ; 
X ISL). ((, lens, jiartly overlapped above bv the iirotiudiiig chitin(*us 
plate /. 

February 3, 1903. 

Howard Saunders, Esq., F.L.S., Vice-President, 
in the Chair. 

The Secretary read the following report on the additions to the 
Society's Menagei'ie during the month of Januaiy 1903 : — 

The registered additions to the Society's Menagerie during the 
mouth of Januaiy 1903 were 58 in number. Of these 15 were 
acquired by presentation and 9 by purchase, and 34 were i'ecei\-ed 
on deposit. The total number of dej)artures during the same 
period, l)y death and removals, was 120. 

The following papers were read : — 

1. Notes on the Hair-Slope of four Typical Mammals. 
By Walter Kidd, F.Z.S. 

[Received November 27, 1902.] 

The Otter. Domestic Dog, Ox, and Horse have been selected 
for consideration as showing very different arrangements of tlieir 
hairy coverings, and as affording by their environments and habits 
the probable explanation of the differences found. Two CarniNores 
and two Ungulates are thus compared and contrasteil. 

(1) In the Otter, taken as a type of the long-liodied hairy 
mannnal with very short limbs, the liair presents an unbi'okeii 
slope from the snout to the tip of tlic tiil. On the liead and 

80 DR. W. KIBD ON THE [Feb. 3, 

trunk it passes from cephalic to caudal, and on the limbs from 
proximal to distal exti'emities in a pei-fectly tiniform manner. 
This ai-rangement obtains in a great number of long-bodied or 
primitive forms, such as rodents, smaller carnivores, insectivores, 
marsupials, lemurs and lower monkeys. As to the hair-slope in 
the Otter, nothing further requires notice, and the type may be 
looked upon as that of the primitive hairy mammal. 

(2) When a Carnivore of a difFei-ent form, such as a short-bodied 
close-haired Domestic Dog, is investigated, a few departures fiom 
the primitive type of hair-slope are found. The Dog is taken as 
representing a more highly developed carnivore form, and it 
presents, as do all Canidfe possessing sufficiently short hair : — ■ 

(i) A whorl situated at the extremity of the snout with a 
feathered arrangement proceeding from it towards 
the orbits, 
(ii) Reversed bilateral area of hair on the pectoral region, 
(iii) Reversed bilateral area of hair on the ventral surface of 

the abdomen in the inguinal region. 
(iv) Bilateral symmetrical whorl in the gluteal region lying 
exactly over the tubera ischii, and, proceeding towards 
this spot, a reversed area of hair on the extensor 
aspect of the thigh. 
(v) Revei'sed area of hair on the extensor surface of the 
These have been fully described, and explanations of their 
setiology put forward, in our Proceedings \ 

The Domestic Dog thus presents an advance in several directions 
upon the primitive arrangement of hair in the Otter. 

(3) The Domestic Ox shows certain peculiarities and departures, 
both from the primitive and specialized Carnivore type : — 

(i) In the frontal region a whorl and a feathering which 
proceeds from it towards the level of the external ears, 
terminating hei'e in a crest. 
(ii) Short longitudinal crest or mane, commencing as a tuft 

at the level of the horns, 
(iii) This crest or mane is interrupted abruptly in its normal 
backward course at about the middle of the neck by a 
ridge. At this point there is produced a meeting of 
two streams of hair pi'oceeding in opposite directions, 
and the forward or " abnormal " stream is found to 
commence at a whorl which lies in the median plane 
at about the middle of the dorsal region. 
(iv) From the whoi'l situated on the dorsal region the slope 
of hair resumes its normal direction and passes to the 
tail, on Avhich it is so ai'ranged that a central longi- 
tudinal crest is usually found, and it terminates in a 
scanty tuft of hair. 

1 Proc. Zool Soc. 1902, vol. ii. pp. 148, 149, 150. 


The Ox presents no pectoi-al whorl as do the Dofj and Ilcn-se, tlie 
dewlap in this region being covered with perfectly smooth and 
uniformly arranged hair. 

These are the only four constant peculiarities of hair-slope, but 
occasionally the flank and post-humeral regions, and extensor 
siTrface of the thigh, show small whorls and featherings connected 
apparently with the action of the panniculus carnosus muscle. 

(4) The Horse is more specialized in the matter of hair-slope 
than any other animal except man. 
It shows constantly — • 

(i) Frontal whorl, feathering, and crest. 

(ii) Tuft between the external ears, 
(iii) Longitudinal crest or niiine. 

(iv) Strong efiicient tail with bushy hair, reaching to the 

(v) Inguinal whoil, feathering, and crest, 
(vi) Pectoral whorl, feathei'ing, and crest. 

Frequently but not constantly : — 

(vii) Post-humeral whorl, feathering, and crest, 
(viii) Cervical whoi'ls, featherings, and crests in several 
(ix) Gluteal whorl, feathering, and crest, 
(x) Tuft on lateral aspect of the abdomen. 

In passing from the simple type found on the Otter to that of 
the Horse, a very significant series of changes is thus displayed. 

Such facts as are here referi-ed to have little or no intrinsic 
interest or importance, but the phenomena of Nature, small and 
great, demand explanation in accordance with the methods of 
science, and it is impossible to ignore the peculiarities of hair- 
slope which have been taken here as typical of a vastly greater 
number in other animal forms. Any other interest they may 
have is subordinate to their relation to the problems of heredity. 
The four groups of facts suggest apparently two explanations as 
to their (etiology. The first and most obvious is that some of them 
ai'e adaptive modifications of value to the animal ; the second, 
that others are produced by its habits ; and it is not difficult to 
distinguish these two classes in the four typical forms chosen for 

1. In the Otter the uniform trend of hair requires no other 
explanation than that this arrangement of hair ofters the least 
possible resistance to mo^■ements in the water and in burrows. 

2. In the case of the Domestic Dog the departures from a primi- 
tive type can hardly be ascribed to anything else than to use or 
habit : they are adapted hy the habits of the animal, not for its 

Proc. Zool. Sor._1903, Vol. I. No. VI. 6 

82 DR. W. KIDD ON THE [Feb. 3^ 

3. The Domestic Ox presents cei-tain points of interest intimately 
associated with adaptive modifications, but many of the phenomena 
are evidently not so associated. Thus, along the median line of 
the dorsal aspect of the Ox is seen a tuft, crest or mane, whorl, 
and long efficient tail, the length of the last being such as to 
reach beyond the dorsal whorl. The large hairy external ears, 
which can be flapped backwards so as nearly to reach the middle 
line, may also be included. These may be looked upon as adaptive 
modifications, existing for the purpose of defending the animal 
against injuiious insects; and a few observations on several oxen 
and horses have been made as to the importance and fi'equency of 
these, and as to the use of the panniculus carnosus, even in this 
temperate climate. 

In moderate summer weather and an exposed wind-swept 
situation, the number of occasions on which certain oxen and 
horses flapped backwards and forwards their ears, corriTgated their 
skins by the action of the panniculus, and flicked their tails 
on to their backs was observed and noted with the following 
results : — 

Tails flicked by oxen at the rate of 348, 468, 504, 540, 720, 

780, 1082 times in an hour. 
Ears flapped by oxen at the rate of 684, 816, 840 times in an 

Panniculus acted in oxen at the rate of 984 times in an hour. 
Tail flicked by horse at the rate of 1108 times in an hour. 

A rough idea may thus be gained as to the importance of certain 
of these mechanisms for defence against injurious insects in hot 
countries, if in a temperate climate and exposed situation so 
frequent a use is made of them. 

4. The Horse also presents in the median plane several similar 
modifications, as a tuft, mane, and long bushy tail, which, when not 
docked, reaches almost exactly to the spot where the mane termi- 
nates. The panniculus in the Horse is much moi-e active in that 
part of the flank and in the forequai'ters where the tail does not 
i-each than elsewhere. 

It remains only to point out the distinction maintained here 
between modifications of hair -arrangement which are themselves 
adaptive, as in the Otter, and others which are, so to speak, by- 
products of habits of the animals exhibiting them, such as whorls, 
featherings, and crests. The only meaning properly assigned to 
adaptive modifications is that such modifications are adapted for 
the needs or comfoi't of an animal. When they ai'e adapted hy 
the habits of an animal, and have no thinkable relation to its 
needs, they must be classed stiictly as non-adaptive phenomena. 

It should be further stated that all the phenomena i-eferred to 
are congenital, and would seem to have an intimate connection 
with the problems of heredity. 


List I. 
Otter No modifications of primitive type of liair-slope. 

Dog 2 whorls itc. 

3 reversed areas of hair. 

Ox 2 whoilsctc. 

Tuft and small mane. 
Crest and tuft on tail. 

Horse ... 3 whorls i-c. ^ 

Tuft. \ ,, , . 

nr - Constant. 


Bushy tail. J 

4 whorls etc. Occasional. 

List II. 


Ox .... 


T, J Phenomena of 

JJepartiires Tr • 

J" It ■ -L- Hair-arranue- 

froin Prinntive I , 7 .■' 

■' ,r . , tinent, adaptive or 

•^ non-adaptive. 


All adaptive. 
All non-adaptive. 


I faint. I. 

4 constant, several I non-adaptive, 
occasional. 1 3 adaptive. 


fi con.stant, 4 oc-! 3 non-adaptive, 
casional. j 3 adaptive. 


Homogeneous. , Simple. 

More varied. More varied. 

Still more varied. Still more varied 

Most varied of 
the ibur t^pes. 

Complex, com- 
binini; tlieliahiti 
of its wild lift 
witli those of 

List III. 

Observations as to use of E.\:ternal Ears, Panniculus 
Carnosu.s, and Tail in Ox and Horse. 

OxEX flapped external ears at the rate of 684, 810, 840 times 

an hour. 

In OxEX panniculus acted at the i-ate of 984 times an houi\ 

OxEX flicked tails at the rate of 348, 468, 504, 540, 

720, 780, 1082 
times an hour. 

Horse flicked tail at the rate of 11 08 times an hour. 

84 CATT. F. WALL OK THE [Feb. 3, 

2. A Prodromns o£ the Snakes hitherto recorded from China^ 
Japan, and the Loo Choo Islands ; -with some Notes. 
Bj Captain F. Wall, Indian Medical Service.^ 

[Received December 15, 1902.] 

"Whilst attached to the China Expeditionary Forces from 
1900-1902, I had opportunities of examining the Snakes pre- 
served in the three Museums in China, and also others in private 
collections. My notes on these, together with those on specimens 
obtained myself, form the subject of this paper, which I have 
arranged so as to form a complete prodromus of the species 
hitherto recorded from the countries above specified. 

In spite of the large aggregate of specimens I examined, it is 
perhaps worthy of remark that I failed to discover one species 
new to science'", and this only serves to show how extensively and 
thoroughly this branch of natural history has been worked out. 

In the City Hall Museum in Hongkong, out of about one 
hundred specimens from the territory above mentioned, I found 
many misnamed and others unidentified. I was informed that 
during a typhoon some years previously a large case, containing 
specimens, was blown over and, the contents wrecked. Out of 
the debris labels were recovered as far as possible and replaced, 
but some were evidently incorrectly reattached and others were 
destroyed. This circumstance may render the accuracy of some 
of the records open to question. That this collection is far from 
representative is evidenced by the fact that during five and a half 
months' residence in this Port I obtained six species which were 
not to be found in the Hongkong Museum. 

In the Museum in Shanghai I found about fifty specimens 
which were for the most part old, and concerning which there 
was practically no information regarding their habitat. 

In the Museum belonging to the Jesuit Fathers at Siccawei, 
near Shanghai, I examined some two or three hundred specimens, 
but these, again, furnished practical!}^ no record of their habitat ; 
however, the late Pere Heude informed me that they had all been 
collected in the Yangtse Yalley, and he mentioned Ning-ko-foo 
(which I find is on the southern bank of the Yangtse River) as 
being the most northern limit. He could not define limits either 
to the south or west. 

As regards habitat, where I do not cite the authority, so far as 
the territory this makes reference to is concerned, it is to be under- 
stood that examples exist in the British Museum Collection at 
South Kensington. 

I have only made reference to points in these specimens which 
do not absolutely agree with the desci'iptions to be found in 

1 Communicated by G. A. Boulengee, F.R.S., V.P.Z.S. 

2 The species referred to hereafter as Dipsadomorphits kraepeUm, No. 65, which 
I had hoped to have the honour of describing for tlie first time, was recorded by 
Stejneger shortly after I began to prepare this paper . 

1903.] SNAKES OF CniXA AXD JAPAN', 85 

Mr. Boulenger's work ' Catalogue of .Snakes in the British 
Museum,' 1893-1896, and 1 have also adopted the nomenclature 
of that woi-k. 

I am indeVjted to Mr. Boulenger for much valuable infoi-mation 
and assistance in drawing up tliis synopsis. My tlianks are also 
due to Mr. F. Bowley, Hongkong, Dr. Barchet, Shangliai, and 
the late Pere Heude, 8iccawei, for their courtesy in granting me 
access to the collections in the Museums under their super- 
intendence, as well as to Mr. A. Owston, Yokohama, and 
Mr. Armsti'ong, Hongkong, for placing their specimens at my 

Part I. — List of Chinese OniiDiA, 


1. TvPHLOPS LiNEATUS. Maliiy Peninsula and Archipelago, 
Hongkong ? 

2. TvPHLors BRAJiiNUS. Southei'u Asia from Ai'ahia to Malay 
Archipelago, South China, Formosa, Hongkong, Hainan, Mau- 
ritius, Comoro Islands, Madagascai-, Cape of Good Hope. — I 
obtained one small specimen three and a half inches long on the 
Peak in Hongkong Island. 


3. Python reticulatus. Burma, Indo-China,, Siam, Malay 
Peninsula and Archipelago, S. China, Hongkong {City Hall 
Mus.). — I saw three specimens, one in the City Hall Museum, 
Hongkong, labelled from that island, one in the Shanghai 
Museum, habitat unrecorded ; and one in the Siccawei Museum, 
consisting of a head only. From the information given by the 
late Pere Heude concerning this collection, I must include this 
species among the Chinese fauna. In all the specimens there are 
three rows of priefrontals, and the lower labials from the 2nd to 
4th and from 13th to 18th are pitted. The frontal is divided 
longitudinally in two of the specimens. A^entrals 315-319. 
Subcaudals 92-100. 

4. Python molurus. Southern Asia from India to China, 
Hainan, Hongkong {City Hall Mas.), Formosa {Sivinhue).— 
The Chinese name given me was " Hiang zo," or " aromatic 
snake," perhaps in reference to their cooked flesh, which is highly 
esteemed by these people. In Formosa it is called " Uang ' 


5. AcROCHORDus JAVANicus. Malay Peninsula, Java, New 
Guinea, Hongkong {City Hall Mvs. No. 293). 

6. PoLYODONTOPnis coLLARis. Himalayas as far west as Simla, 
Assam, Arrakan, Upper Burma, S. China, Hongkong. — I ob- 
tained one specimen on Hongkong Island. There is one anterior 

^6 CAPT. K. WALL ON THE [Feb. 3, 

temporal on both sides, wbicli comes in contact with the eighth 
labial only ; this latter is unusually high and is probably the 
result of a confluence with the normal inferior anterior temporal. 
The first lower labials do not meet behind the symphysis. Belly 

7. Tropidonotus savinhonis. Formosa. 

8. Tropidonotus nuchalis. China. 

9. Tropidonotus vibakari. Manchuria, Japan, Formosa. 

10. Tropidonotus piscator. Southern Asia from India to 
China, Hainan {Herz & City Hall Mus.), Hongkong {Hallow. & 
City Hall Mus.), Formosa \Stejneger). — Apparently common in 
the extreme south. I obtained three specimens from the main- 
land opposite Hongkong, and saw one in Mr. Armsti'ong's col- 
lection which he assured me he procured from Hongkong Island. 
I found only two specimens in the large Siccawei Collection. In 
one specimen (froan Kowloon, opposite Hongkong) there ai-e five 
postoculars on one side, in another four on both sides. In one 
there are eight upper labials, with the fourth only touching the 
eye on both sides. 

11. Tropidoxotus annularis. China, Formosa. — This must 
be a very common snake in the Yangtse A^alley, judging from the 
large numbei- of specimens in the Siccawei Museum, but in the 
extreme south it appears to be rare or absent. There are two 
prfeoculars on both sides in one specimen, five postoculars on 
both sides in one specimen, and two postoculars on one side 
in one specimen. Ventrals 132-164. Subcaudals 70 in one 

12. Tropidonotus tigrinus. Siam, Cochin China, China, 
Hainan, Hongkong ? Manchuria, Corea, Japan. — Called " Yeh- 
chi-po" by the Chin :se, signifying pheasant's neck, also " Ch'ing- 
ch'ang-chung {Moll.). Apparently as common as T. annularis 
in the Yangtse Valley, from the number of specimens at Siccawei. 
One specimen in Mr. Armstrong's collection was, he informed me, 
captured on Hongkong Island ; however, it was not labelled, and 
his collection contained some species he had procured in Japan. 

13. Tropidonotus stolatus. Southern Asia from India to 
China, Hainan, Hongkong, Formosa, Chusan Archipelago, Philip- 
pines. — I obtained one specimen from the mainland opposite 
Hongkong. The labials on one side were nine in number, and 
the fourth, fifth, and sixth touched the eye. I saw no specimen 
in the Shanghai or Siccaw^ei Collections. 

14. Tropidonotus subminiatus. Eastern Himalayas, Assam, 
Burma, Malay Peniasula and Archipelago, S. China, Hongkong. 

15. Tropidonotus chrysargus. Eastern Himalayas, Assam, 
Burma, Malay Peninsula and Archipelago, S. China, Hainan. 


16. Tropidoxotus balteatus (Cope), Proc. Ac. Phil;ul. 1894, 
p. 426. H:tiiian. 

17. Tropidoxotits craspedogaster (Blyr.), 1^. Z. S. Loml. 181)9, 
p. 163, pi. xvii. fig. 1. China. 

18. Tropiuoxotus percarixatus (Blgr.), P. Z. 8. Load. 1899, 
J). 163, J)!, xvii. fig. 2. China. — I saw six specimens in tlie 
Siccawei Collection which exactly fit the description of Mr. Bou • 
lenger's specimen except in the following details: — Length of 
internasals sometimes eipials pi-jefrontals. Postocnlars and sub- 
oculars : the numbers of shields in contact with the back of tlie 
eye, intervening between the supraocular above and the fifth 
labial, are four in two specimens on both sides, four in one 
specimen on one side, five in two specimens on both sides, fi\e 
in two specimens on one side, six in one specimen on one side. 
Labials eight with third and fourth touching the eye on one 
side, nine with fifth only touching the eye on one side. Ventrals 
and subcaudiils 140 + 73, 139+ ?, 139-^73, 138+ ?, 138 + 72 (tip 
slightly docked). 

19. Tapixophis LATOucnn (Blgr.), P.Z. S. Lond. 1899, p. 164, 
pi. xviii. figs. 1-1 c. China. 

20. PsEUDoxEXODOX MACROPS. Him:d'\yas, Khasi Hills, Hills 
in Biu'ma, Yunnan [de Scahra^), and S.W. China. 


22. Opisthotropis axdersoxii. Hongkong, — I obtained five 
specimens, all from Hongkong Island. They were all captured in 
a swamp near the Sanatorium on the Peak, whilst being drained 
during the campaign against malai-ial mosquitoes. One was dug 
up at a depth of about 2 feet below the surface. They accord 
with Mr. Boulenger's description except in the following parti- 
culars : — Labials are inconsistent in ai-rangement. There are nine 
in one specimen on one side, eight in all the rest. The fourth 
and fifth touch the eye in one specimen on one si<lc, the fifth 
only touches the eye in two cases on both sides, and in one case 
on one side, no labials touch the eye in two specimens on one 
side. Pmeoculars (including the suboculars of Boulenger) are 
two in two specimens on both sides, two in two specimens on one 
side, one in one specimen on one side, and three in one .specimen 
on one side. The anterior chin-shields are in contact with five 
lower labials in two specimens on both sides, and in one specimen 
on one side, with four in one specimen on both sides, and in 
one specimen on one side. Ventrals and subcaudals 167 + 60, 
149 + 53, 164 + 59, 165 + 59. Colour uniform dull olive-bluish 
above, lower half of ultimate row and belly yellow. Lower labials 
and throat-scales with dull bluish mottling. Sparse mottling 
beneath tail. 

1 JJull. Mus. ir. N. Paris, 1«>7, iii. V- 215. 

88 CAPT. F. WALL ON THE [Feb. 3, 

23. Trirhinopholis styani (Blgi\), P. Z. S. Lond. 1899, p. 164, 
pi. xviii. figs. 2 & 2 a. China. 

24. AcHALii^us RUPESCENS. Hoiigkong. — I obtained four 
specimens in Hongkong Island, found in the low vegetation on 
the slopes of the Peak. In one specimen both anterior temporals 
touch the eye on both sides. Scales somewhat irregular, 23-25 
in mid-body. Yentrals and subcaudals 150 + 56, 158 + 58, 158 + 
61, 154 + 58. Colour uniform olive-brown above, slightly darker 
vertebrally, and with iridescence in reflected light. Head same 
colour above, merging to chestnut on temporal regions. 

25. AcHALiNus BRACCONiERi. S. China. — I think that this 
species will have to be united with A . spinalis ; the differences 
between the two shown by Mr. Boulenger ^ are considerable but 
not constant, being shared by individuals of both supposed species. 
For instance, in all the four specimens of A. spinalis that I have 
examined (one of which I presented to the British Museum, 
which has been seen and identified as such by Mr. Boulenger) 
the scales are 23, and not 21. The relative length of the sutures 
between the internasal and prsefrontal has, in my opinion, little 
or no weight. The specimen of A. spinalis 1 sent to the British 
Museum has the internasal suture about two-thirds the length of 
the praefrontal ; and in more than one specimen of hracconieri in 
the British Museum a similar condition exists. The specimen 
figured by Stejneger ^ from Japan which he calls spinalis is more 
like Mr. Boulenger's hracconieri, but has 23 scales. 

26. ACHALINUS SPINALIS. China, Japan.- — I saw three speci- 
mens in Mr. Owston's collection from Mount Fuji, Japan, and 
one I found in the Siccawei Collection. In all there is a large 
shield on the postero-lateral region of the parietals similar to 
those described under ^4. h-acconieri, and, like them, separated by 
one scale in the median line. Ventrals and subcaudals 154 + 58, 
165 + 44, 165 + 48, 170 + 44. 

27. Lycodon aulicus. Southern Asia from India to Malay 
Archipelago, Philippines, Formosa {City Hall Mus.), Hongkong? 
{Boettger ^), S. China ? viz. from Amoy {Steindachner *). 

38. Lycodon fasciat^us. W. Yunnan [Anderson), Assam, 

29. Lycodon subcinctus. Malay Peninsula and Archipelago, 
Sumatra, Borneo, Java, Timor, Hongkong, Philippines. — There 
are two specimens in the City Hall Museum from Hongkong and 
one from Timor. I also examined a fourth, which Mr. Armstrong 

1 Cat. Snakes Brit. Mus. vol. i. pp. 308 & 309. 

2 Ann. Zool. .Tapon. ii. p. 29. 

3 The specimen alluded to by Boettger (Mat. herp. Faun, von China, 1888, p. 84) 
in the City Hall .Museiiin is no longer in the collection; the only specimen of this 
species in that In^stitution is i'rom Formosa. 

'' Eeise der Novai-a, Rept., Wien, 1869, p. 74. 


informed me he had captured in the filter-beds near Bowen Road 
on Hongkong Island. In two specimens the third, fourth, fifth, 
and sixth supralabials touch the eye on both sides. Ventrals 
192-209 \ 

30. DixoDOX RUFozoNATUS. China, Hainan, Formosa, Chusan, 
Corea, Tsu Shima, Japan, Loo Choo Islands. — This is a very 
common snake about Shanghai, whei-e I encountered it frequently. 
I picked up several and allowed them to ci-awl about my arms 
without their exhibiting the least malice or alarm, and they 
made little attempt to escape in the first instance. Riding my 
bicycle one night I saw one crossing the road ; I dismounted, 
flashed my lamp on the snake, and, while holding my machine 
Avith my light hand, captured it easily with my left. It made no 
attempt at escape, though cover was within a yard or two. It 
Avas full-grown and not des(]unmating. My servant caught one 
in camp one night just outside my hut ; it encircled the man's 
leg but did not bite. It contained a large toad {Bufo vulgaris). 
I found a large number preserved in the Siccawei Museum. I 
found one in Mr. Owston's collection obtained from Japnn, and 
four othei-s procui'ed fi'om Ishigaki Island in the Loo Choo group. 
I noted the following : — Internasals half or less than half the 
length of pr^efrontals. Loreal, in Japanese and Loo Choo speci- 
mens, does not touch the eye in all (five) ; in Chinese does not 
touch the eye in five, touches eye in eleven. Postoculars three 
on both sides in one specimen, three on one side in one specimen. 
Temporals one on one side in one specimen. Labials normal in 
all. Anterior chin-shields in contact with four lower labials in 
one specimen on both sides, with six lower labials on one side 
in two specimens. Ventrals in Japanese and Loo Choo speci- 
mens 180-190; in Chinese specimens 192-209. Subcaudals iia 
Japanese and Loo Choo specimens 76-87 ; in Chinese specimens 
64-76. Colour : there appear to be two very distinct varieties. 
All the Chinese conform to the following description : — Alternate 
bars of jet-ljlaek and coiul-i'ed (white in old spu-it-specimens) 
doi'Siilly, breaking into a coai'se mottling on the flanks. The black 
bars involve two or three scales in the length of the snake, and the 
red one scale or slightly moi-e. Thei'e ai-e 53-74 black bars on 
the body and 18-24 on the tail, the first is broadest and forms a 
chevron on the nape. Head black, fading to wdiitish on labials ; 
sutures on crown coral-red. A light temporal streak usually. 
Belly whitish, with some lateral mottling. The Japanese and 
Loo Choo specimens agree : — Alternate darkish-bi-own and dirty 
whitish (perhaps red when fresh) bars doi'sally, breaking up into 
a mottling laterally. The bi'own bars involve four or five scales 
(more quite anteriorly), the light one scale. Thei-e are 24-33 
brown bai's on body, 15-18 on tail. Crown of head broAvn, fading 

1 Giinther records also Li/codnn (OpJtilcs) alLofiiscus from Formosa (Ann. Mafr. 
Nat. Hist. (4) vol. i. 180S, p. 420), but jjives no dcscni'tion nor authority for 
recording it. 

90 CAPT. F. WALL ON THE [Feb. 3, 

to whitish on labials. A light temporal streak. Belly whitish, 
with or without sparse lateiul mottling, except beneath tail 
where this is abundant. 

31. DiNODON SEPTENTRiONALis. China, Formosa, Himalayas? 

32. Zaocys DHUMNADES. S. China, Chusan. — A very common 
snake about Shanghai and evidently throughout the Yangtse 
Valley, for there are many specimens in the Siccawei Museum. 
I have encountered it frequently, and consider its generic name 
most apt, as it is veiy active, swift, and clever to elude capture. 
I saw as many as four in a day's ramble in the spring, when 
batrachians wei-e clamouiing in the water engaged on matrimonial 
matters. The snakes wei-e each coiled vip on the banks close to 
the water, and in low vegetation, awaiting the excursion of some 
unwary individual. One I saw coiled up similarly two yards or 
so away from two toads {Biifo vulgaris), Avhose curious behaviour 
it was that first attracted my attention. In spite of every cai-e 
my presence was always detected by the snake before I was 
aware of its presence, and it immediately made off in great haste 
for the water, and disappeared among the i-oots of the aquatic 
vegetation. On one occasion in the summer I watched one for 
some time in a strip of grass in the open, myself unobserved ; 
and it was most interesting to notice the method and care 
with which it beat the patch of gi-ass like a hai-rier, prying into 
every recess in the ground or tussock that might harbour some 
possible prey. I have seen Chinese jugglers wdth this snake in 
their stock in trade, and I believe that it is this species that 
is even now occasionally met with in houses in the heart of 
the town of Shanghai. On October 3rd, 1901, I captured a 
young one recently hatched in camp which closely resembled the 
adult in colouring. Loreal : in one specimen there are two 
superposed shields on one side. Temporals : a single anterior 
in one specimen on one side. Anterior chin-shields in contact 
with four lower labials on both sides in one specimen. Ventrals 
190-199. Subcaudals 96-119. 

33. Zamenis korp^os. Sikkim Himalayas, Assam, Burma, 
Siam, Malay Peninsula, Sumatra, Java, W. Yunnan, S. China, 
Hainan, Hongkong, Formosa {City Hall Mus.). — I obtained one 
specimen on the mainland opposite Hongkong. There are two 
specimens in the Siccawei Museum. Loreal single on both sides 
in one specimen, three on one side in one specimen. Subcaudals 

34. Zamenis mucosus. Southern Asia from Transcaspia to 
Malay Peninsula, Java, S. China, Hainan {City Hall Mas. <& Herz), 
Hongkong, Formosa, Chusan {Cantor d' Gilnther). — I obtained 
two examples from Stonecutter's Island in Hongkong Harbour, 
and three from the mainland opposite Hongkong. There are two 
specimens in the Siccawei Museum. Labials nine, the fifth and 
sixth touching the eye in one specimen on one side. 

1903.] SXAFvES OF C'lilXA AXD JAPAN'. 91 

35. Zamexis snxALis. Siani, H;iin:in, China, Coroa, IMonijolia. 
This snake is e\'idently veiy common in the Valley, as 
there are ten specimens in the Siccawei IVIuseum. Concerning 
one which I obtained from Huangtsun, N. China, my donor wrote 
me that he had found it in his verandah, it having dropped, lie 
believed, from the i-oof abovit 7 feet above. He discovei'e(l it in the 
act of devouiing a lizai'd [Crecko suhpalmatas). It had seized the 
gecko by the body and was encircling it in its coils. A little later, 
Avhen the captive had exhausted its futile struggles, the snake 
seized it by the snout and began to swallow it. Frontal three- 
fourths to foui'-fiftlis parietals. Loreals : two (anteiior and 
posterior) on both sides in one specimen. Subocular absent in one 
specimen on one side. Praeoculars : the upper touches the frontal 
in two specimens on both sides. Tempoiuls : two anterior in all 
specimens. Labials : eight, with the fourth and fiftli touching 
the eye in two specimens only, the normal ariungement being 
nine, with the fifth and sixth touching the eye. Antei'ior chin- 
shields in contact with four lower lahials in two specimens on both 
sides. Ventrals 179-207. Subcaudals 84-100. 

36. Coluber porphyraceus. E. Himalayas, Hills in Assam, 
Burma, Malay Peninsula, Siuuatra, Yunnan {Anderson). 

37. Coluber maxdarixus. Chusan, China (N.W. Fokien, 
Blgr. ; Prov. Chihli, Mijll.). I saw one specimen in the Siccawei 
Collection. It is called by the Chinese " 'Hua-tai-tsze," meaning 
variegated girdle {Mull.). 

38. Coluber rufodorsatus. E. Siberia, China, Chusan, For- 
mosa, Hainan. — Evidently a very common snake. There are fi\e 
in the Shanghai Museum, and a large number at Siccawei. 
Frontal often greater than distance to end of snout. Anterior 
chin-shields in contact with four infralabials in one specimen out 
of fifteen examined. 

39. Coluber diox'E. S. Russia, Transcaucasia, Temperate Asia, 
China, Hainan, Japan [Blgr.). — Evidently a veiy counnon snake 
in the Yangtse Valley, as the Siccawei Collection contains a large 
number of specimens. I obtained two from an officer in Ching- 
wang-tao, N. China, who told me the snake was common there. 
One specimen contained the brood of some small bird, four in 
number, one of which was still partly env-eloped in its shell. 
Frontal often greater than its distance to end of snout, sometimes 
equalling parietals. Pra-oculars : three on one side in one speci- 
men. Postoculars : one on both sides in one specimen out of 
sixteen examined. Labials nine, with fourth and fifth touching 
the eye on one side in one specimen ; nine with fifth and sixth 
touching the eye on one side in two sjiecimens only. Anteiior 
chin-shield often greater than posterioi', in contact Avith six 
'infralabials on both sides in one specimen, and on one side in one 
s})ecimeu. Scales in mid-body 23 in three specimens, 25 in ten, 

92 CAPT. F. WALL ON THE ' [Feb. 3, 

and 27 in one specimen. This species is called " Huang-ch'ang- 
ch'ing " by the Chinese [IfdlL). 

40. Coluber t^niurus. Sikkim, Cochin China, Siam, Malay 
Peninsula, Borneo, Sumatra, China, Formosa {/Senckenberg Mus., 
Boettg.), Chusan, Manchui-ia. — This snake is common in the 
Yangtse Valley. I encountered one near Shanghai, which, however, 
I failed to captui'e as it took to water and disappeared in the 
submerged vegetation, Loreals confluent with prfefrontals on both 
sides in one specimen. Temporals : single on both sides in one 
specimen, and on one side in one specimen. Labials : nine with 
the foui-th and fifth touching the eye in one specimen on both sides. 
Ventrals 226-242. 

41. Coluber schrenckii. Amoorland, Corea, N. Japan. 

42. Coluber phyllophis. China. — I saw seven specimens in 
the Siccawei Museum, the largest measuring 8 feet 1 inch. There 
is also a stuffed specimen in the Shanghai Museum. Scales 21 
in mid-body in one specimen. Ventrals 221 in one specimen. 

43. Coluber davidi. China {Scmvage). 

44. Coluber mcellendorffii. China {Ilerz d; Broeclcelmann). 

45. Coluber melanurus. Burma, Malay Peninsula, Borneo, 
Sumatra, Java, S. China.— I saw one specimen in the City Hall 
Museum from Java, Subcaudals 88. 

46. Coluber radiatus. E. Himalayas, Bengal, Assam, Burma, 
Cochin China, Malay Peninsula, Sumatra, Java, S. China {Miill., 
fferz, Moll., Kaufm., Broeckl.), Hongkong {City Hall Mus.). — 
There are five specimens in the City Hall Museum from Hong- 
kong. I also obtained one specimen from that island which 
contained four blind and callow offspring, the brood of some small 
mammal, pi'obably a rat \ 

47. Dendrophis pictus. S. Asia from India to Malay Archi- 
pelago, Cliina {Giinth.), Hongkong, Philippines. — One specimen 
of this snake in spirit was given me by a lady, who assured me 
she had obtained it on Hongkong Island. 

48. SiMOTES purpurasoens. Siam, Cochin China, Malay 
Peninsula, Sumatiu, Borneo, Java, S. China ~. 

49. SiMOTES CYCLURUS. Bengal, Assam, Burma, Siam, Cochin 
China, Malay Peninsula, Sumatra, S. China. 

50. SiMOTES FORMOSANUS. S. China, Formosa, 

51. SiMOTES viOLACEUS. Bengal, Assam, Burma, Camboja, 

-^ Coluber climacoplwrus vel SlapJiis virgatus, described by Boettger (Mat. hei'p. 
Faun, von China, 1888, p. 72) from Corea, is, 1 think, an error. The specimSn 
alluded to is evidently Coluber schrenckii. 

2 pam^ril & Bibrpii, 'Erp^fcologie,' p. 632. 

1903.] sxAKES orcnixA AND jArAx. 93 

B. Cliina, Hainan, Hongkong. — I saw one specimen in tlie City 
Hall Musenni, and obtained two myself on the mainland opposite 
Hongkong. One was caught in grass on the hillside and suffeved 
itself to be handled without I'etaliiition. The other was caught on 
a shi'ub poised over a flower. Frontal : length greater than dis- 
tance to end of snout in three specimens. Labials : six with the 
third and fourth touching the eye on one side, seven with the 
fourth and fifth toucliing the eye on the other side in one speci- 
men. Nasals semidi\ided on both sides in one specimen. 

52. SiMOTES CHixEXSis. China, Hainan. — Rostral : visible 
portion seen from above equals distance to frontal in two 
specimens. Labials seven, with the fourth only touching the eye 
on both sides in one sj)ecimen. Subcaudals 50 in one and 51 in 
another specimen. I saw two specimens in the Sicca wei Museum, 
and obtained one myself fi'om Kiangyin on the southern bank of 
the Yangtse River, to the noi-th-west of Shanghai. 

53. SiMOTES vAiLLANTi. China (Sauvage). 

54. Ablabes major. China, Hongkong [Gunth., IlaUow., City 
JIall Mits.), Formosa, Chusan Archipelago. — Evidently fairly 
common in the Yangtse Valley. I saw one specimen of this snake, 
belonging to ]\Tr. Styan, in Shanghai, and many others in the 
Siccawei Collection. Thei'e are thi'ee in the City Hall Museum, 
two of which ai'e fi'om Formosa, the other of nnceitain habitat. 
I saw one specimen in Mr. Armstrong's collection in Hongkong 
obtained from that isLuid, whei'e, as he told me, it is not 
uncommon. Labials : in one specimen seven, with the third 
and fourth touching the eye on one side, Subcaudals 89 in one 
specimen, and 90 in two others. 

55. Ablabes dori/E. Kachin Hills, Assam, China. — I saw one 
small specimen 8 or 9 inches long in the Siccawei Collection. 
Yentrals 160 ? Subcaudals 85 ? It agrees in every particular 
with Mr. Boulengei-'s desci'iption. This species has not been 
previously recorded from China, 

56. Calamaria pavimextata. Burma, Siam, Cochin CliiiL-i, 
Malay Peninsula, Java, S, China {Moll.), Riu Kiu Archipelago 
[Stejneger '). 

57. Calamaria berezowski, S. China {Gilnth. ^). 

58. Calamaria septextrioxalls. China, Chusan Archipelago, 
Hongkong. — I saw two specimens in the Sicwxwei Collection, both 
quite typical. 

59. SPAXiornoLLS souliei. Yunnan {de Scabra ^). 

^ Stejneger (I'roc. Biol. Soc. Wasli. xiv. p. 191) ilcserihes a snake .is new to scieiioe 
under the name of Calamaria pfejfcri, which, in my opinion, is u specimen of 
C pitvimentdla. 

■ Ann Ae. St. Petersb. 1896. p. 205, pi. i. fig-, a. 

^ Bvill. .Mas. N. II. V:\v\%, iii. 18S)7, p. 211!. 

94 CAPT. F. WALL ON THE [Feb. 3, 

60. HypsiRHiNA PLUMBEA. Burma, Inrlo-China, Malay Penin- 
sula and Archipelago, S. China, Hainan, Hongkong, Formosa.— I 
saw two specimens in the City Hall Museum, one labelled China 
and one Hongkong, and I obtained four myself on the mainland 
opposite Hongkong. All my specimens were caught in or about 
a sluggish stream near our camp at Kowloon. Frontal : length 
greater than distance to end of snout in six specimens. Anterior 
chin-shields in contact with six infralabials in two specimens on 
one side. I noticed that in some specimens the lateral scales 
about the anal region had minute central tubercles, reminding 
one of the condition in Aspidura cojyii and A. trachyprocta. 

61. Hyfsirhixa enhydris. India, Ceylon, Burma, Siam, 
Cochin China, Malay Peninsula, Borneo, Celebes, S. China, 

62. Hypsiriiina bennetti. China, Hainan (Ilerz). 

63. Hypsirhina chinensis. Siam, China, Hainan, Hongkong 
{Steindachner '). 

64. HoMALOPSis BUCCATA. Bengal ?, Burma, Indo-China, Malay 
Peninsula, Sumatiu, Borneo, Java, Hongkong {licdloto. <k City 
Hall Mus.). — I examined the specimen in the City Hall Museum 
which is labelled Hongkong. 

65. DiPSADOMORPHUS KRAEPELixi : Boiga JiraepeXini Stejneger 
(Proc. Biol. Soc. Wash. 1901, xv. p. 15). Formosa. — I found one 
old specimen in the City Hall Museum labelled Formosa. This 
specimen almost exactly accords with Stejneger's description, the 
only points of diflterence being :^ — Temporals are six on one side. 
Labials ten on one side. Anterioi- chin-shields in contact with 
five lower labials on both sides. Yentials 244. Subcaudals 140. 

Malay Peninsula, Sumatra, Java, Celebes, S. China, Hongkong. — 
A common snake in the south of China,. I obtained six from 
Hongkong Island, one from Stonecutter's Island in that harbour, 
and one from the mainland opposite Hongkong. There are 
several specimens in the City Hall Museum, and I saw others in 
spirit and in captivity belonging to Mr. Armstrong. 

and Assam Hills, Burma, Indo-China, Malay Peninsula and 
Archipelago, Formosa. 

68. Chrysopelea orxata. Ceylon, Hills of S. India, Bengal, 
Assam, Burma,, Indo-China, Malay Peninsula and Archipelago, 
S. China, Hongkong ? ' — I found one specimen in the Siccawei 

1 Reise der Novara, Kept., Wieii, 1869, p. 68. 

- The specimen referred to by Boettger (Mat. lieryi. Fauna von China, p. 142) 
is no longer in the City Hall Museum. 


IVIusenm and one in the tSlifinghai Musenin. Suliciii'lals 1J4 in 
one specimen. 

69. AiPYSURUS AXxrLAxrs. Loyalty Islands, Loo Choos, 8ca,s 
around Formosa {iS'iejneger)\ 

70. Hydrus platurus. Oliok, lied Sea, Indian Ocean, Straits 
of Malacca, Tropical and Snl)tro]iical Pacific fi'om Loo Choo to 
Australia and New Zealand, Malay Arcliipelago to Central 
America. — I saw two specimens in the City Hall Museum fi'om 
Hongkong and Formosa, and one in the Shanghai Museum. Also 
two or three specimens in Mr. Owston's collection in Yokohama, 
obtaineil t'l-om the Japanese shoi-es or the Loo Choos. All })elong to 
variety E of the British Museum Catalogue of Snakes. Fi'ontal : 
length less than distance to end of snout in one specimen, less than 
parietals in two. Labials nine on one side in two specimens ; none 
bordering the eye on one side in two sjjecimens. Anal tetrafid. 

7L AcALYP'iDriiis PEROXii, Western Tropical Pacific. 

72. HYDiiornis fasciatus. Coasts of India to China and Xew 

73. Hydrophis GRACILIS, Coasts from Persia to China. 
Hainan [Iler.z). 

74. Hydrophis melaxocephalus. Indian Ocean, Pescadoi'es, 
{Stejneger)", Loo Choos. 

75. Hydrophis obscurus. Bay of Bengal to China. Canton 
{Peters) \ 

76. DiSTiRA STOKESii. Mekran Coast to Chinese Sea, and Korth 
Coast of Australia. 

77. DiSTiRA ORXATA. Coasts of Asia from mouth of Persian 
Gulf to Loo Choos, Kew Guinea, and N. Australia. — Tliis snake 
evidently is connnon in the Loo Choos, as I saw more than twenty 
specimens from that region in Mr. Owston's collection. 

78. DiSTiRA SUBCIXCTA. Indian Ocean, Japan, Loo Choos 
{Siejueyer) '. 

' Sti'jiK'.troi- (.Jnnrn. Sc. (V)ll. Tokyo, xii. 18!)S-lfl()0, p. 223) <k'scrilK>s as !i uow 
spet'ies uiultT i\w name of l<l>ui/docepIialus ijimce certain specimens, tlie ilesci-iption of 
wliich I have consulted ]\Ir. Boulen.uer aliout ; and he is of ojiininn lliat these will he 
found to \w ^\vm\\vu^ o\' A i fiiisHnni ail inihi f iig. The enlar<red vertehralsand ]n-esenco 
of four iirn'trontals, found in these s]iecinu'ns, were also present in some specimens 
I examined lielon.uin.n- to .Air. Owston in Yokohama, which [ re.ii'arded at the time as 
anniilatnst. An examination of the specimens in the British Museum shows that a 
certain slijrht cnlar;;cniunt of the vertehrals is present at some sj'ots, hut no specimen 
has four iira'frontals. '{"he wci'^ht of .such an opinion compels me to modify the views 
J had formed on the suhject. 

- .J. Coll. .laiian. xii. p. 221 CMicroccplinlophis mihiiiorc[>h(tI ti!<). 

» Monatsh. Herlin. Akad. 1872. p. 8o9 {H. dhttlcmn). 

' Steineser (I'roc. Hiol. Soc. Wash. xiv. ]>. 1!)1. l'.)01) deseriltes certain specimens 
oli'tained from the I'iu Kin Seas as Dixlim i>n<-)ii(i/is, which he considiM-s a new 
si)ecie^. Kmm hi- ili-,-ri|it ion I am oti.pininn that these helonj;- to Jj. suhcinrfd. 

96 CAPT. F. WALL ON THE [Feb, 3, 

79. DiSTiRA BRUGMAXsii. Persian Gulf to Chinese Sea. 
Hainan [jBoettg.) \ 

80. DiSTiRA CYANOCiNCTA. Persian Gulf to Chinese Seas, 
Japan, Papuasia. — I saw one specimen of this snake in the City 
Hall Museum labelled Hydrus viajor from Hongkong, and one in 
the Shanghai Museum. One conforms to type A, and the other 
to type C of the British Museum Catalogue. 

81. DiSTiRA viPERiNA. Persian Gulf to Chinese Sea. 

82. Enhydris hardwickii. Bay of Bengal to Chinese Sea and 
Coast of New Guinea. — There are four specimens in the City Hall 
Museum : two, labelled Hydrus major, are from Manila, and two 
from Bangkok. There is one specimen in the Shanghai Museum 
with very markedly spinose tubei'cles on the median six ventral 
rows of scales. These rows are also enlarg'ed. Labials eight in 
two specimens, in one on both sides and in the other on one side 

83. Platurus laticaudatus ^. Bay of Bengal to Chinese Sea, 
Loo Choos, New Guinea, and Western South Pacific Ocean. 
There are three specimens in the City Hall Mviseum labelled 
Formosa, and I saw several specimens in Mr. Owston's collection 
obtained from the Loo Choo Islands. Ventrals 232-246. Sub- 
caudals 32-46. 

84. Platurus colubrinus. Bay of Bengal to Chinese Sea, 
Western South Pacific Ocean. — I examined two specimens, both 
in the City Hall Museum, one from Penang and the other of 
uncertain habitat. 

85. BuNGARUS FASCIATUS. Southern Asia from Bengal to 
China, Hongkong {City Hall Mus.). — I obtained one specimen 
from the mainland opposite Hongkong, and I saw two others in 
the City Hall Museum, one from Hongkong and the other from 
the mainland opposite. 

86. BuNGARUS CAXDiDus. Southern Asia from India to China, 
Hainan, Hongkong {City Hall Mus.), Formosa. — I saw two speci- 
mens in the Shanghai Museum, one in the Siccawei Collection, 
and one in the possession of Mr. Styan in Shanghai. There are 
four specimens in the City Hall Museum, one from Hainan and 
three from Hongkong. All these specimens are of variety B of 
the British Museum Catalogue, viz. midticincta. Postoculars : 
the normal lower shield confluent with the fourth labial on both 
sides in one specimen. Labials six, with the second and thh-d 

1 Under the name of Sijdro^Jhis cyanncinctus Boettger (Mat. herp. Faun, von 
China, 1888, p. 88) describes three specimens of what I consider Dis^ira hriigmansii , 
collected b}' Herz in Hainan. 

^ The species described by Boulenger (Cat. Snulces Brit. Mus. iii. p. 309) as Platurus 
muelJeri I do not believe to be valid, and I think the specimens on which it is 
based will prove to be F. laticaudatus. The only definite point he mentions to 
characterize it is the presence of a median ventral keel in the posterior half of the 
body, and I have found this peculiarity in at least three specimens of what I consider 
uudoubted P. laticaudatus. 


touching the eye on one side in one specimen, and occasioned by a 
confluence between tlie first and second labials. Anterior chin- 
shields in contact with four infralabials on both sides in tliree out 
of four specimens examined. 

87. ISTaia tripudians. Southern Asia from Ti-anscaspia to 
China, Hainan, Hongkong [Jlalloio., Steindach., City Hall Mus.), 
Formosa, Chusan, Philippines. — In the Siccawei Museum 1 saw 
seven specimens, and in the Shanghai Museum three. There are 
fifteen in the City Hall Museum, of which eleven are from Hong- 
kong, two from Hainan, and two from China. Of the ten 
specimens I examined three have scales in twenty-one rows in 
the middle of the body, and three in nineteen. In the remainder 
I have failed to record the number. Nine have more or less 
distinct (some very well defined) buft" or pale yellowish cross-bands 
dorsally. These numbered from 13-21 on the body, and 5-8 on the 
tail. They involve one or two rows of scales along the length of 
the snake, and are most conspicuous in the posterior third or so 
of the body. The intervals involve fiom ten to twelve scales, and 
are sometimes sepai'ated from cross-bars by a blackish line. Hood 
marked with modified black ocellus. Belly yellow, with one or two 
plumbeous bands ventrally, or mottled to a variable extent with 
black. In one specimen the whole belly uniform black. One 
specimen is evidently a Sputatiix. This I found in the Siccawei 
Collection. Scales over hood 21, mid-body 17. Ventrals 177. 
Subcaudals 45. No hood-marks. Nearly unifoi-m black dorsally, 
with no suspicion of cross-bars. One very broad plumbeous ventral 
band involves from the 9th to the 47th shield, and then breaks up 
into a mottling and disappears. 

88. Naia bungarus. Southern Asia from India to S. China, 
Hongkong {City Hall Mus.), Philippines. — A newly-hatched ex- 
ample is preserved in the City Hall Museum, habitat Hongkong. 
I found the head of a large specimen in the Shanghai Museum ; 
and whilst I was in Hongkong a gentleman encountered one ou 
the mainland opposite, which he killed and bi'ought home. I 
examined and identified it, and estimated it at between seven and 
eight feet in length. 

89. Callophis macclellandii. Nepal, Sikkim, Assam, Burma, 
S. China, Formosa, 

A M B L Y C E P 11 A L I D .«. 

90. Amblycephalus moellendorffii. Tenasserim, Siam, Cochin 
China, S. China, Hongkong, Hainan. — This is a common snake 
in the island of Hongkong. I collected ten myself, all of which 
were found in low jungle on the slopes of the Peak. I saw three 
specimens in the City Hall Museum. Internasals form a suture 
with the loreal in all specimens. Postoculars and suboculars : 
usually one long semilunar shield extends from the supraoculars 
behind, skirting the eye to a point about halfway up its anterior 
aspect. Sometimes this scale is divided so as to form a small 

Proc. Zool. Soc— 1903, Vol. I. No. YII. 7 

98 CAPT. p. WALL OX THE • [Feb. 3, 

postocular above this shield. Temporals : the antero-superior 
extends back as far as the extreme border of the parietals or 
further. Labials : six in two specimens on both sides. Sub- 
caudals 36-50. First infralabials meet in four, fail to meet in 
one, not recorded in eight. 

91. PsEUDOPAREAS VAGUS. Hongkong (Jan). 


92. Ancistrodon acutus. China. — Called by the Chinese 
" Oo-woo-shay," or five-pace snake, owing to the reputed rapidly 
fatal eifects of its bite. I saw one specimen in the City Hall 
Museum labelled A. blomhoffii, habitat Szechuen. There are also 
five specimens in the Sicca wei Collection. Two are labelled Ou- 
Yuen\ and one of these dated 1882 (i. e. six years before Giinther 
first described it). One is labelled Chowtong" 1883, and another 
Kien-te^ 1882. The fifth consists of a dried skin. In talking 
to the late P^re Heude about this creature, he told me it is not 
uncommon in hilly parts of the Yangtse Yalley, and is feared by 
the natives more than a leopard. He narrated an experience of 
one of the Jesuit Fathers who heard frantic cries for help issuing 
from some dense jungle he was riding through. On dismounting, 
and proceeding to the spot, he found a Chinaman being pursued 
by one of the snakes, which he attacked and killed. Rostral : as 
far as I could ascertain there is only one shield beneath the dermal 
appendage, which, however, is frequently bent over, causing a 
fuiTow which makes the shield appear divided into two. Labials : 
six in one specimen on both sides, and in another on one side. 
Subcaudals 56 in one specimen, with Ist-llth entire and rest 
divided ; 53 in another, with lst-9th entire and rest divided ; 58 
in another, with lst-28th entire and rest divided ; 57 in another, 
with lst-12th entire and rest divided. 

93. Ancistrddon blomhoffii. E. Siberia, Mongolia, China, 
Hainan, Formosa, Japan, Loo Choos, Siam. — Mollendoi-flf says it 
is called by the Chinese " Fei-shang-ts'ao," snake which flits on 
the grass, and " Ch'i-ts'un-tsze," or seven-inch snake. It is very 
common in the Yangtse Yalley. There are at least three specimens 
in the City Hall Museum from Japan. There are five in the 
Shanghai Museum, and a very large number in the Siccawei 
Collection. In Mr. Owston's collection I saw several obtained 
from Japan and the Loo Choos. I obtained five myself about 
Shanghai in our camp. Two of these, which I found close together 
on October 16th, 1901, appeared to be newly hatched. On the 
4th October, 1901, my servant informed me that there were several 
snakes lying together dead close to the camp. I investigated the 
matter, and found an adult of this species with the remains of 

1 Perhaps Tao-yuan in Province Hunau ? 
" Perhaps Chao-tung- in North Yunnan ? 
5 In Province An-Hui. 


twelve ynimg PCiittered ai'ouud her. In spite of every endeavour, 
I failed to tr:ice the author of this butchery. It is .«jid luck tliat 
such oppoi'tunities as these for oliservino- hahits I'ai'ely fall to the 
lot of tliose who specially seek them. Residents ahout Shanghai 
occasionally lose dogs under circumstances suggesting a poi.sonoua 
bite, and I think these fatalities are frequently due to this little 
snake. One specimen I cjiptvu-ed had the penis hifid on both sides. 
I kept young on two or thi-ee occasions, but could not find suitable 
food with which to tempt them, and finally I'esorted to feeding 
them with boluses of raw meat, a treatment they displayed their 
distaste for l)y repeatedly disgorging the morsels some time later. 
I observed that, when molested and excited, all my specimens 
vibrated the tail, but I could never get one actually to sti'ike ; 
however all the specimens I had alive were veiy small. Labials 
seven, the third only touching the eye ; eight in one specimen on 
both sides. 

94. Lachesis jerdoxii. Ivhasi Hills, Assam, Thibet ; Upper 
Yangtse, China. 

95. Lachesis mucrosquamatus. Kaga Hills, Assam ; Formosa. 

96. Lachesis cramixeus. South-eastern Asia, China, Hainan 
{Herz (I- Cit]/ Hall Mas.), Hongkong, Formosa. — This is a veiy 
common snake in Hongkong, whci'e I procui'ed three specimens. 
There are also twelve in the City Hall Museum, obtained locally. 
In one of these the skin of the neck had been perforated and 
rent by the beak of a bird which it is seen in the act of devouring. 
The distension of the mouth and neck is enormous, vso that the bird 
appears as if it were being thrust through the rent instead of into 
the gullet. Thei'e is only one specimen in the Shanghai Museum, 
and only one in the Siccawei Collection. The lesidents of Hong- 
kong call this the Bamboo-snake, which is a very good name for it 
since it is almost always to be found in the foliage of bamboo 
vegetation. I am told too that the Anglo-Indians in Assam have 
also given this name to the snake. 

Part II. — List of Japaxese axd Loo Choo Islaxds Ophidia. 

C O L U B R I D --E. 

1. Tropidoxotus vibakari. Japan, Formosa, Manchuria. — I 
found one in the City Hall Museum from Nagasaki, and saw six 
others in Mr. Owston's collection, all from IVlount Fuji. Ventrals 
152 in one specimen. Subcaudals 80 in one specimen; second to 
fifth entire in one specimen. 

2. Tropidoxotus tigrixus. Siam, Cochin China, China, Hainan, 
Hongkong ?, Manchuria. — In my opinion this is the commonest 
snake in Japan. I frequently met with it, and on one occa'^ion 
captured five in a couple of hours. It is called by the Japanese 
"yamakagashi," " uwabami," '-orochi," "'ja," all of which terms, I 


100 CAPT. F. WALL ON THE [Feb. 3, 

am informed, signify " the largest variety " ; also " tora-no-kuchi- 
nawa," or "tiger-snake"; also " miza kuchinawa," or "water- 
snake"; also "atsuki kuchinawa," or "thick brown snake." (See 
also No. 12 of Chinese list.) 

3. Tropidonotus prteri. Loo Choos. — I saw one specimen in 
Mr. Owston's collection from ISTawa in the Loo Choos. 

4. AcHALiNUS SPINALIS. China, Japan. (See No. 26 of Chinese 

5. DiNODON RUFOZONATUS. China, Hainan, Formosa, Chiisan, 
Corea, Tsn Shima, Japan, Loo Choos. (See No. 30 of Chinese list.) 


7. DiNODON JAPONicus. Japan. — I found one specimen on the 
Southern Island (Kiu Siu), and saw another in Mr. Owston's 
collection. Yentrals 209. Subcaudals 75 ? and 77. 


9. Coluber dione. S. Russia, Transcaucasia, Temperate Asia, 
China, Hainan, Japan (Blgr.). (See No. 39 of Chinese list.) 

10. Coluber schrenckii. Amoorland, Corea, N. Japan. 

11. Coluber conspicillatus. Japan, Corea. — I saw one 
specimen in the City Hall Museum labelled Nagasaki and another 
in Mr. Armstrong's collection captured in Japan. Called by 
Japanese "jimuguri" ("earth borer") and " kawara kuchinawa" 
(" dry-river-bed snake "). 

12. Coluber climacophorus. Japan. — A very common snake 
in Japan, where it is called "aodaisho," "nezumi tori" (or " i-at- 
catcher"), "sato meguri" (or "village idler"), and " mugiwara hebi." 
I frequently encountered it. I saw and captured one reclining 
on a stone parapet in one of the Nikko temples. I discovered 
another in thick jungle in the act of swallowing a half-grown 
leveret, ti-uly an enormous meal, since the mammal was more 
than twice the weight and girth of the reptile. I thought on 
all occasions it was far less wary and active than other nearly 
allied snakes of a similar size with which I am familiar, such, for 
instance, as Zamenis 7)111008118 and Z. korros, Coluber radiatus, 
Zaocys dhumnades, &c. Temporals, in one specimen, three on 
both sides. 

13. Coluber quadrivirgatus. Japan, Corea. — This species is 
almost if not quite as common in Japan as Tro^ndonottis tigrimis. 
I scarcely went a day in the coiintry without seeing one, and 
often I saw three or four. It is called by the Japanese " shima 
hebi" (or "striped variety"), "karasu hebi" (or "black snake"), 
" sukuro hebi" (or " medium black snake "), " mugiwara hebi " (or 
" wheat straw snake"), and " kiiro kuchinawa." Subocular absent 
in one specimen on one side. Labials eight, with third, fourth, 
and fifth touching the eye on one side in one specimen. 


14. Coluber schmackeri. Loo Choos. 

15. Ablabes semicarixatus. Japan, Loo Choos. 

16. Ablabes hermin.e. Loo Choos. — I saw two specimens in 
Mr. Owston's collection from the Loo Choos. Ventrals 162-163. 
Subcaudals 47-51. 

17. Calamaria pavimextata. Burma, Siam, Cochin Cliina, 
Malay Peninsula, Java, S. China (Mull.), Riu Kiu Archipelago 
(Stejneger). (See No. 56 of Chinese List and footnote.) 

18. Hydrus platurus. Coasts of Asia from Red Sea to Loo 
Choos, Australia, New Zealand, Malay Archipelago to Central 
America. — Called " umi hebi" or "sea snake" by Japanese. (See 
No. 70 in Chinese list.) 

19. Hydrophis melaxocephalus. Indian Ocean, Pescadores 
{Stejneger), Loo Choos. — This is evidently a common snake about 
the Loo Choos, since I saw upwards of twenty specimens in 
IMr. Owston's collection from this locality. Postoculars one or two. 
Temporals : one anterior usually, sometimes two. Scales two 
head-lengths from snout 23-27, mid-body 31-35. Yeutrals 305- 
343. Anal tetrafid. (See No. 74 of Chinese list.) 

20. DiSTiRA cyaxocixcta. Persian Gulf to Chinese Seas, 
Japan, Papuasia. (See No. 80 of Chinese list.) 

21. DiSTiRA ORXATA. Coasts of Asia from mouth of Persian 
Gulf to Loo Choos, New Guinea, Northern Australia. (See 
No. 77 of Chinese list.) 

22. DiSTiRA subcixcta. Indian Ocean, Japan, Loo Choos 
{Stejneger). (See No. 78 of Chinese list and footnote.) 

23. Aipysurus axxulatus. Loyalty Islands, Loo Choos, Seas 
around Formosa {Stejneger). (See No. 67 of Chinese list and 

24. Platurus laticaudatus. Bay of Bengal to Chinese Sea, 
Loo Choos, New Guinea, Western South Pacific Ocean. (See 
No. 83 of Chinese list.) 

25. Platurus colubrixus. Bay of Bengal to Chinese Sea, 
Western South Pacific Ocean. (See No. 84 of Chinese list.) 

26. Platurus schistorhyxchus. Western Pacific Ocean, Loo 

27. Hemibuxgarus japoxicus. Loo Choos, Japan ? 


28. AxcisTRODOx ixxESMEDius. Japan, Mongolia, Ea^ern 
Siberia, Central Asia. 


29. Ancistrodon blomhoffii. Eastern Siberia, Mongolia, 
China, Hainan, Formosa, Japan, Loo Choos, Siam. — Called by 
the Japanese " mamushi," " hami," " knchibami," "hiraguchi." 
All synonymous terms for the " beautifvilly marked variety." 

30. Lachesis okinavensis. Loo Choos. — Called by the Japanese 
" habu." 

3L Lachesis flavoviridis. Loo Choos. — Also called "habu" 
by the Japanese. 

32. Lachesis mucrosquamatus. Naga Hills, Assam ; Formosa. 

33. Lachesis luteus. Loo Choos. 

3. Note on the Wild Sheep of the Kopet-Dagh. 
By R. Lydekker. 

[Eeceived December 4, 1903.] 
(Text-figure 10.) 

Tlirough the generosity of Mr. St. George Littledale the collec- 
tion of the British Museum has recently been eniiched by a very 
fine skull, with the horns (text-fig. 10, p. 103), of a ram of the Wild 
Sheep of the Kopet-Dagh i-ange, which forms the boundary between 
Turkestan and Noi-thern Pei-sia to the eastward of the Caucasus. 
Mr. Littledale also brought home the skin of the same animal, 
bvit it unfortunately was so badly injured by vermin that it had 
to be destroyed. The skull is the finest of the series obtained 
during the trip. 

The Kopet-Dagh sheep was named Ovis arkal in 1857 by 
Blasius, and is evidently allied to the Urial, with the Punjab 
race of which I have indeed proposed to identify it \ At that 
time I had, however, never seen an adult skull ; and Mr. Little- 
dale's specimen indicates the right of this sheep to rank as a 
distinct race of Uriah It wall be remembered that the Punjab 
race of the Urial {0. vignei cycloceros), as exemplified at any rate 
by specimens from Peshawer and Afghanistan in the British 
Museum, difiers from the typical 0. vignei of Astor and Ladak 
by the much greater prominence of the two front angles of the 
horns, which are often raised into nodose beads, between which 
the front surface of the horn is depressed and carries bold and 
widely separated ti'ansverse ridges. 

In the Kopet-Dagh Urial this prominence of the front angles 
of the horns is still more pronounced, though the beading is 
somewhat less conspicuous. Moreover, the fi'ont surface of the 
horn is unusually broad and flattened, with the transverse 

1 ' Wild Oxen, Sheep, and Goats,' p. 173. 




♦wi'inkles very low and indistinct. The It'ngth of the horn is 
33 inches along the inner front angle, with a basal circnniference 
of 11 inches, a basal width of 3 inches, and a ba.sal depth of 
4 inches. 

The last two dimensions are considerably greater than in a 
skull of the Kelat Urial measured by Mr. Hume, in which the 
length along the curve is 35| inches. 

Text-tiij. 10. 

Front view of skull and lionis of adult rain of the Kopet-Dagh Urial. 
5 nat. size. 

The Kopet Dagh Urial decidedly appears to be a distinct form, 
connected with the typical Ocis vignei by means of the Punjab 
race of that species. On these grounds I regard it as a local race 
I'ather than a species ; its name will accordingly be 0. vignei ai-kal 
(or perhaps arcal). The suggestion of M. Dauvergne \ that this 
sheep is identical with the Kelat Urial, is not borne out by a 
comparison of the present specimen with a skull of that form in 
the British Museum, in which the angles of the horns are much 
rounded ofi\ This leads me to think that the Kelat Urial 
{0. vignei blanfordi) is, after all, distinct from the Punjab 

1 See p. 131 of my book on 'Game of Europe, N.W. Asia, and America.' 

104 MR. P. W. BASSETT-SMITH OlST [Feb. 3, 

4. On new Parasitic Copepoda from Zanzibar and East Africa, 
collected by Mr. Cyril Crossland, B.A., B.Sc. By Staff- 
Surgeon P. W. Bassett-Smith, R.N., F.Z.S. 

[Received December 4, 1902.] 
(Text-figures 11 & 12.) 

Mr. Cyril Crossland, in his recent examination of the marine 
fauna of Zanzibar and British East Afi'ica, obtained several 
specimens of parasitic and semiparasitic Copepods, three of which 
he has been kind enough to allow me to examine. 

These curiously deformed and often grotesquely-shaped animals 
are frequently found attached to the gills, &e., or to the surface of 
fish and other marine animals. 

A large number from the former which ai-e now described from 
a variety of different hosts, and from wide geographical areas, 
I enumerated in Proc. Zool. Soc. 1899, p. 438 ; to this paper 
I have appended a list of addenda, which I have drawn up with 
the kind assistance of Mr. E. Bergrotti, of Tammerfers, and 

Not only fish bvxt a number of other marine animals are un- 
doubtedly infested with these parasites, though at present little 
information concerning them has been collected ; the specimens of 
Mr. Crossland are therefore particularly interesting. 

Gerstacker, in Bronn's ' KJass. unci Ordn. des Thier- Reich s,' 
1866-79, Crustacea, vol. v. Copepoda, p. 774, mentions five genera 
found on Nudi branch Mollusca : Doridicola Lyd., Eolidicola Sars, 
belonging to the family Ergasilidae ; Ai-totrogus Boeck, to the family 
Ascomyzontidse ; and Siolanchnoiroinis Hanc. and Ismailia Bergh, 
to the family Chrondi'acanthidas. Also nine genera fi'om various 
Vermes, p. 773. 

Of the three specimens of Mr. Crossland, two were taken from 
the kidneys of species of Pleurobranchids (not determined) and 
one from the skin of a Sipunculid (^Aspidosiphon). 

As they were only single specimens it was impossible to dissect 
them, and therefore the descriptions are necessarily incomplete. 
The first two evidently belong to the family Chondracanthidre, 
but do not fall in with the descriptions of any known genus ; they 
appear to be most nearly related to the genvis Splanchnotropus of 
Hancock (Trans. Linn. Soc. vol. xxiv. pp. 51, 55), two species of 
Avhich he describes, *S'. gracilis and S. brevipes, taken from Nudi- 
branchs ; the present specimens differ from them, however, in the 
complete absence of antennae and articulate limbs, and in having 
the external ovaries elongated and the eggs arranged in single 

I have therefore provisionally placed them in a new genus, 
Cho7ulroca7-pus, following closely after S2^lconchnotro2n(,s Hanc. and 
IHocus Fabr. 

Chondrocarpus, gen. nov. 

$ . Cephalothorax coriaceous, elongated, with four lateral short 




lobe-like processes on either side ; abdomen one-fourth length of 
whole, biiU'tieulate, tapering ; no distinct antennae or thoracic 
limbs ; mouth placed on under surface, with minute maxillte ; 
external ovaries as elongated filiform sacs containing ova in a 
single series, 
d. Pigmy. 

Chondrocarpus reticulosus, sp. n. (Text-fig. 11, A-G.) 

llah. Zi;\\\z\\rAV : from large Pleurobi'anchid, 

Length 12 mm., breailtli 4 nnn. 

2 . Oephalothorax indistinctly segmented, elongated, tapering, 
dorsally convex, laterally produced into 4 rounded truncated 
processes ; the 4th pair being most widely separated and the 
smallest ; the whole having a peculiar I'eticulate appearance from 
the network of ramifying tubules; anterior extremity rounded; 
no visible antenna; ; mouth as a papilla placed between the 1st 
pair of processes ; upper lip triangular ; oidy one pair of maxillae 
and mandibles (?) could be made out, each terminating in minute 
claws. No thoi-acic limbs; there is a short genital segment, or^ 
ring, from which spring the long filiform ovaries. Abdomen in- 
distinctly biarticulate, tapering, without caudal plates or seta\ 

<S . Pigmy. One was seen attached to the last abdominal 
segment but was partially hidden, the bifid ai'ticulate caudal 
extremity only being visible. 

Text-fig. 11. 

Chondrocarpus reticulosus ? , pren. & sp. ii. A. Dorsal surface. B. Lateral view. 
C. First segment and mouth-organs. D. One of tlic lobes sliowing reticu- 
late appearance. E. Second maxilliped. 1<\ Abdomen showing fixed j. 
G. Posterior extremity of (J. H. C/iondrocarjius sp. $ : dor»al surface 
(specimen incomplete). 

CnoxDRocARPus sp. (Text-fig. 11, II.) 

A specimen of a second species of this genus was taken from a 

106 MR. P. W. BASSETT-SMITH ON [Feb. 3, 

Pleurobranchid ; it was much broken both at the anterior and 
posterior extremities. It differs from G. reticidosiijS in not having 
the peculiar reticulate appearance and in having a pair of lateral 
lobes on either side of the genital segment. 

The third specimen also appears to be new, and belongs to the 
family Dichelesthiidfe, the animal resembling most nearly the 
genus Enterocola of Van Beneden (Bull. Acad. Roy. de Belg. 
torn. ix. 2nd ser. p. 151), found by him in the respiratory cavity of 
" Aphidium Jicus," than any other form I have been able to find 

Mr. Crosstand's specimen appears to be much more degenerate 
from its parasitic habits ; the ai'ticulate limbs are excessively 
small, difficult to make out, and the ova are carried in long spiral 
thread-like processes in a single series as in the Caligidse, and 
not in dilated sacs. Unfortunately there was only this single 
female specimen for examination. I would pi'ovisionally ci-eate 
for it a new genus " Ventriculina," giving the specific name of 
" crosslandi" in recognition of the collector. 

Ventriculina, gen. nov. 

Head small, rounded ; neck indistinct ; 3 thoracic segments, 
the first amalgamated with the head ; genital segments lobed, 
equal in breadth with the thoracic ; no dorsal plates ; abdomen 
short, biarticulate ; external ovaries spiral, ova uniserial ; first 
antennse 4-jointed, simple, second antennae 3-jointed ; maxillipeds 
very small , 

Three pairs of minvite thoi'acic limbs, first biramose, second and 
third uniramose. 

Ventriculina crosslandi, sp. n. (Text-fig. 12, p. 107.) 

Hah. Zanzibar : from a Sipunculid. 

Total length 4 mm. Colour white. 

Head small, rounded in front, broadest behind, from under 
which pi'oject the anterior antennae. Thoracic segments three, 
the first united with the head, the genital segments trilobed ; the 
whole forming an oblong body without lamellar plates, showing 
five distinct rounded lateral lobes, the last pair being slightly 
wider and more acute. Dorsal surface convex, marked by five 
distinct grooves showing the position of the union of the segment^. 

Abdomen biarticidate, narrow, one-sixth the total length, the 
last joint terminating in two small caudal plates provided with a 
marginal fringe of short bi'istles. External ovaries long, spiral, 
spi'inging fi'om papillse at the angle of the genital segments and 
abdomen. Ova large, arranged uniserially as in the Caligidse. 

Anterior antennae 4-jointed, non-se''jOse, the first joint being 
the longest and broadest, rising from the underside of the head 
just in front of the rnouth ; second, third, and fourth joints 
progressively decreasing in size. 




Posterior antennre S-joiuted, rising just in front of the upper 
lip ; at the distal end and anterior border of the first and second 
joints are two short setje, the third terminating in two bristles, 
the anterior being very long. 

Mouth and appendages placed rather far back ; the labrum is 
tiiangular, projecting backwards ; labium simple, I'ounded. I was 
able to make out only 2 pairs of maxillipeds ; the first very small, 
biarticulate, terminating in tw^o short hairs : the second uncinate, 
with large globose basal joint, to wdiich was articulated a sharp 
curved claw. 

Text-fig. 12. 

Vciitriculhia crosslandi $, gen. & sp. n. A. Ventral surface, X 10. B. Dorsal 
surface. C. Ventral surtacc much enlarged, showing articulate appendages. 
D. Posterior antenna;. 

Only three pau-s of thoracic limbs present : the first rising f i-om 
the posterior under sui-face of the cephalic segment, minute, 
biramose, each i-amus terminating in a single bi'istle ; second and 
thii'd paii's uniiumose, made up of two ai'ticulations, the distal 
tei-minating in two small bristles. 

(S not known. 

Addenda to Systematic Enumeration of Sjxicies of Parasitic 
Copepoda found on Fish (Proc. Zool. Soc. 1891), p. 438). 


1. Eucanthns viarchesettii Yalle, Atti Mus. Civ. Trieste, vii. 

p. 245 (1885). On Jlotella tricirrata. 

2. Ergasilas centrarchidaricm Wright, Proc. Canad. Inst. (2) i. 

p. 243 (1883). 

3. Eryasilus biuncinatus Gadd. Meddelanden af Societas pro 

Fauna et Flora Fennica, xxvii. pp. 181 182 (1901). On 
Gastrostens acnleatns. 


4. Bomolochus onosi T. Scott, 20th Ann. Rep. Fish. Board of 

Scotland, p. 289, pi. xiii. figs. 19-22 (1902). On Onos 
musiehts and O710S cimbrius, Firth of Forth. 

5. Bomolochus zeugopteri T. Scott, loc. cit. p. 290, pi. xiii. figs. 23-25. 

On Zeugojiterus punctatus. 


P. Caligus pacificus Gissler, Amer. ISTat. p. 886 (1883). On 

7. Caligus labracis T. Scott, I. c. p. 291, pi. xiii. figs. 26-29. On 

Lahriis mixius and L. 'macidatus. 

8. Anchicaligtis nautili Stebbing, Willey's Zool. Res. pt. v. pp. 667- 

670, pi. Ixxi. 

9. Dinematura mtisteli Icevis Hesse, Rev. Sci. Nat. Montpellier, 

(2)ii. pp. 6, 11 (1880). 

10. Cecrops acanthice vulgaris Hesse, Ann. Sci. ISTat. (6) xv. 3, 

p. 26 (1883). 

11. Pandarus carcharii glaucus Hesse, I. c. p. 18. 

12. ,, musteli Icevis Hesse, I. c. p. 23. 

13. ,, spinacis acanthice Hesse, I. c. p. 10. 

14. ,, unicolor Hesse, I. c. p. 20. On Gcdeus vulgaris. 


15. Lernanthropus polyneini Rich. Zool. Anz. iv. 1881, p. 505. 

On Polynevius. 

16. Lernanthropus tetradactylus (probably L. trifolicUus B.-S. 


17. Lernanthropus micro2)terygis Rich. Atti Soc. Tosc. Sci. Nat. 

iv. p. 82 (1884). On Micropteryx dumerili. 

18. Lernanthropus tylosuri Rich. I. c. p. 83. On Tylosurus im- 


19. Kroyeria (Lonchidium) gcdii vidgaris Hesse, Ann. Sci. Nat. 

(6) xvi. 3, p. 2 (1883). 

20. Clavella cluthce T. Scott, l. c. p. 292, pi. xii. figs. 26-31, On 

Ctenolabrus rupestris. 

21. Pagodina (Nemesis) charcharice glauci Hesse, I. c. p. 13. 

22. Eudactylina carcharice glauci Hesse, I. c. p. 11. 

23. ,, musteli Icevis Hesse, I. c. p. 8. 

24. „ squatina ccngeli Hesse, I. c. p. 5. 

25. „ similis T. Scott, I. c. p. 295, pi. xii. figs. 1-19. 
On Raia radiata. 

26. Eudactylina acanthii T. Scott, I. c. p. 296, pi. xiii. figs. 1-9, 

On Squalus acanthias. 

27. Bassettia congri Stebbing, Willey's Zool. Res. pt. v. pp. 671, 

672, pi. Ixx. 


28. Philichthys ficdolce Rich. Zool. Anz. iii. p. 69 (1880). On 

Stromateus jialola. 

29. Philichthys doderleini Rich. Zool. Anz. vi, p. 558 (1883). On 

Labrus turdus. 

1903.] ox THE ORIGIN'Ar. HOME OF THE TIOER. 109 


30. Lerncea ahyssicola Brady, Chall. Rep. viii. p. 137. On Circdias 


31. Lerncea minuta T. Scott, IStli Rep. Fish. Boai'd of Scotland, 

p. 161, pi. vii. fig. 13 (1900). Oa Gobiiis miniUas. 

32. Lermea lumpl T. Scott, 19th ditto, p. 128, pi. vii. fig. 12 

(1901). On C)/clopterus lunipus. 

33. Hfemohaphes amhigims T. Scott, 18tli ditto, p. 1G2, pi. vii, 

fig. 15. On CalUonymus maculatas. 

34. Peraderma petersi Rich. Zool. Anz. iv. 1881, p. 387. On 

Goh'ms huGcatiis. 

35. Peraderma hellottii Rich. Zool. Anz. v. 1882, p. 475. On 

iScopelus henotti. 


36. Chondracanthus hleeheri Rich. Zool. Anz. iv. p. 387 (1881). 

On Chiliiim chlorurus. 

37. Chondracanthus ninnii Rich. Zool. Anz. v. p. 504 (1882). On 


38. Chondracanthus ornatus T. Scott, 20th Ann. Rep. Fish Board 

of Scotland, p. 298, pi. xiii. fig. 34. On CalUonymus 


39. Achtheres sandrce Cladd. Med. af Soc. pro Fauna et Flora 

Fennica, xxvii. (1901). 

40. Lernceopoda extumescens Gadd. I. c. On Coregonus. 

41. Tracheliastes gigas Rich. Zool. Anz. iv. 1881, p. 504. 

42. Charopiims dubius T. Scott, 19th Ann. Rep. Fish Board of 

Scotland, p. 130, pi. vii. fig. 15. On Paia circular is. 

5. On tlie Original Home of the Tiger. 
By Col. C. E. Stewart, G.B., C.M.G., C.I.E.i 

[Received December 6, 1902.] 

The ordinary idea of English people that the Tiger was origin- 
ally an Indian animal, is, I believe, quite a mistake. After 
careful enquiry, I have come to the conclusion that the Tiger is a 
comparatively late intruder into India. 

Firstly, after enquiry, I cnn discover no Sanscrit word for the 
Tiger. If tigers had existed in India in the days when Sansci-it 
was a spoken language, there would he a name in Sanscrit for it, 
while there is only a modern Hindustani name. There is a 
Sanscrit word for Lion, " Singha," which would point to the 
fact that lions were certainly more common than tigers in time 
long past. At present lions are not found in India, except a 
very few, which are strictly preserved in Googerat, one extreme 

' ComniuiiicateJ by Col. Hili, .James, F.Z.S. 


corner of India, tliough I will allow that lions were probably 
commoner tlian they are now in the olden time, though probably 
never veiy numerous. 

I remember, when I first went to India, nearly 50 years ago, a 
lion being killed not very far to the southward of Allahabad, but 
this was even then a rare occurrence. I have studied the 
question of the habitat of lions and tigers in Persia, where I 
resided for a good many years. Lions are found only in the 
very south of Persia, near the Persian Gulf, and Arabia ; while 
tigers are only seen in the very north of Persia, near the Russian 
border, and especially near the Caspian Sea, on the north of 
Persia, and they are moi-e numerous within British territory 
than within the Persian boundary, and tigers are more common 
in Southern Siberia than they are anywhere in Persia. 

Tigers are more numerous in cold countries. They are plen- 
tiful in Oorea, which has a severe winter climate, and still more 
plentiful in the Island of Saghalien, belonging to Russia, and 
further noi'th than Oorea, and which has almost an ai'ctic climate 
in winter. The tiger is mentioned by Marco Polo in his travels, but 
nowhere as an Indian animal, and I very much doubt whether 
tigei'S were found in India at the time Marco Polo visited it. 

In the Sanscrit woi'ks treating of the fighting between Rama 
and Rawun, the Demon King of Ceylon, though many animals 
are mentioned, such as beai-s, monkeys, and several others, I 
have been unable to find any mention of the tiger ; and the tiger 
is not found in the Island of Ceylon, though the leopard is ; nor 
is the tiger found in the larger island of Borneo, which would 
seem to point to its only inhabiting the islands of the Indian 
Archipelago, which it could reach by swimming. Thus it would 
seem that tigers did not exist in India before the time that Ceylon 
was separated from India. Tigers did not exist in the island of 
Singapore until about 1809, when apparently they swam over 
from the mainland. Tigers are such good swimmers that they can 
cross a considerable body of water. I do not think any allusion 
to tigers in India can be found in the Greek historians. I should 
feel much obliged if anyone could find me such a reference. 

In the monuments of the Assyrian Kings, and of the Kings of 
Persia, there are constant references to lion-hunts by those kings, 
but never allusion to a tiger-hunt. Of course there is an existing 
Persian word for tiger, but there is nothing to show that it is at 
all ancient. 

My own idea is that the tiger was originally a purely northern 
animal, which has gradually extended southward. I fancy that 
no allusion to a tiger in India can be traced to a period anterior 
to the early Mahommedan conquerors of India. I should be 
much obliged to anyone who will help me to clear up this 
question. We English have so completely assumed the idea 
that the tiger is an Indian animal, that we have called him the 
Royal Bengal Tiger, though I firmly believe he is as much an 
intruder from the noitli into Bengal as we are ourselves. 

1903.] ox THE roruLATiox of the ixdiax elephant. Ill 

6. On the Mode of Copuliitioii of the Indian Ele]»hant. 
By H. Slade, Conservator of Forests, Maymyo, Burma.^ 

[Received December 15, 1902.] 

Some few months ago I was, on several occasions, al)le to witness 
tame Elephants in tlie act of copulation. This sight has been so 
seldom witnessed by Europeans, and is so variously described l)y 
Burmans, that these remarks, supported V)y a sei'ies of photographs 
deposited with the Society, may be of interest and value. 

As tuskers are usually i-eputed to be shy of copulating before 
eye-witnesses, an account of the manner in which these photo- 
graphs wei-e procured may be interesting. 

I wjxs in camp at the time with one tusker and four female 
Elephants, which were being used regulai-ly for ti'ansport pui'ijoses. 
From the time the tusker was i-epoi-ted to be seeking the company 
of the females he was never let loose to gi'aze, but was kept tied 
up. I had pi'omised the Bui'mese mahouts libei'al i-ewai'ds if they 
would assist me in procuiing some " snap-shots"; and one djiy last 
Febi'uary one of them came to tell me the tuskei- was showing 
undoubted excitement ; he was reported to have been tugging at 
his chain and looking " nastily " at his keeper. There weie, 
however, absolutely no signs of " must " and no exudation of fluid 
fi'om the hole in the temple. 

The tuskei' was said to have shown a prefei'ence for one of the 
females that had calved about 12 months pi-eviously. So I had 
her tethei-ed foi-e and aft in a small glade, and ei'ected my camera 
quite in the open, about 10 yards off. Having focussed her and 
got eveiything I'eady, the tusker was quietly ridden up behind 
the female, and as the mahout slipped ofl", he slowly advanced 
towai'ds hei'. However, this was evidently the WTong female, for 
on the appi'oach of the tusker she showed most unmistaka1)le 
signs of fear by pei'sistently screaming and straining at hei' chain. 
The tusker was therefore secui'ed, and this female marched oft', 
whilst one of the others was caught and tied up in her place. 
The tusker was then again brought out, and immediately pi-oceeded 
to mount the female. 

I had many oppoi-tunities of witnessing the operation, and the 
following is a general desci'iption : — The female, when in season, 
remains perfectly still and quiet, merely signifying her sense of 
the tusker's approach by moving hei- tail slightly to one side and 
gently shifting hei- hind feet a few inches further apart. Tlie 
tusker creeps up behind the female and begins to show signs of 
sexual excitement. He then I'aises his head and lightly places 
his tusks on the female's back, one on either side of her backbone, 
with his trunk lying along the dorsal ridge and reaching to lier 
shoulder. In this position he I'emains a length of time, varying 
with his state of sexual excitement. From my oliservations 1 am 
convinced that when he lisis uniestrieted access to females, and is 

^ Coiniminicatcd Ii}- tlie Seckktaky. 


not therefore greatly excited, he may remain a very considerable 
time in this position : this has probably given rise to the tales 
that are told of the great length of time occupied by the act of 
copulation, which I have heard put at two hours. 

Eventually, however, the time arrives when the tusker proceeds 
to action. First one fore foot, and then the other, is lifted off the 
ground, and slowly swung from side to side across his front, 
exactly as though he were deliberating from which foot to take 
off. Then, levering himself up by his tusks, he places both fore 
feet on the female's flanks, at the same time sinking down on his 
hind legs to an almost sitting position. It is at this moment that 
he shows the most violent sexual excitement. His organ, which 
up till now has been bent back in the form of an arc, with its end 
near the ground, and a few inches only in front of his hind feet, 
is shot forward, and jerked up and down in a most violent manner. 
It is flourished in the air, and often bent into the form of an S 
like the thong of a whip, during which semen may or may not be 
emitted. At one moment the organ appears outside the tusker's 
shoulder, the next it is seen proti-uding out behind his hind legs. 
It is kept in a state of continual agitation, and not for one single 
moment is it still. Eventually it is jerked up straight forward 
and hits the female organ from below. Penetration is then 
immediately effected without further difficulty. The tusker then 
raises himself until his hind legs are quite straight, his organ is 
pushed home, and his fore feet are slid along the female's back 
until they rest on her shoulders. Arrived at this position, he 
begins to work very much after the fashion of a dog, and I 
usually counted from six to eight horizontal motions. Connection 
being then complete, the tusker lifts his tusks ofi" the female's 
back and raises his head into the air. In this position he remains 
for a few seconds, and then slowly withdraws his organ, letting 
himself gently down to the ground in the same way that he 
mounted, and qiiietly moves off. It is then that the female 
shows her only signs of excitement : she trumpets softly as with 
pleasure, thrusts forward her ears and stiffens her tail, her whole 
conduct being indicative of pleasure and pride. The tusker is 
throughout perfectly silent. 

It is noteworthy that at no time does the tusker use his fore legs 
to steady himself or to grip the female as does a dog or a stallion ; 
the feet are invariably kept close together on the top of the female's 
back, and I can quite believe the statement of tlie Burmans that 
it makes no difference whether his feet are fettered or not. 

I was unable to distinguish any sign by which I could tell when 
a female was in season, but three out of the four females in camp 
did come into season during the period of 10 days in February 
during which they were under observation. 

The operation was accurately timed on one occasion, and was 
found to last exactly one minute. 

From my own observations, I am convinced it is the female 
that comes in season, and that until she does so come the tusker 


1/ 1 \.ri-»'' 


Z.D & S.J H. del. 

Eale k Darnels son L*"'^ M. 



will take no notice of her : also that he is rejuly whenever she is. 
I can ofFei' no opinion as to whether the companionship of the 
tusker tends to bi'ing the female into season or not. 

I nuist draw attention to the way in which both the tusker and 
female absolutely ignoi'ed oui' presence. As I have already st;ited, 
the camera was set up within 10 yai-ds of the tethered female, 
quite out in the open witliout any attempt at concealment ; and 
though there were sometimes six or eight spectators walking about 
and talking within this short distance, on no occasion did either 
the tusker or the female take the slightest notice of us or even look 
our way. Dii'cctly the act of copulation was over, the maliout 
called out to the tusker, who, at the word of command, came towards 
him, knelt down, and allowed himself to be mounted and ridden ofi". 

As I have said, the female was always tethei'ed fore and aft 
with a long chain, but this was solely with the object of prevent- 
ing her swinging round and so getting out of focus ; but in no 
single instance did she attempt to do this, nor was the chain once 
needed except, of course, in the case of the first female who was 
not in season. 

7. On the Cffilenterata collected by Mr. C. Crosslaud in 
Zanzibar. — I. Ceratella minima, n. sp. By Sydney J. 
HiCKSON, M.A., F.R.S., F.Z.S., Beyer Professor of 
Zoology in the Owens College, Manchester. 
[Received December 16, 1902.] 
(Plate XIII. ^) 

Our knowledge of the remarkable family of Hydi-ozoa, the 
Cei-atellada^, has been ably summarized in the memoir pul)lished 
by Prof. Spencer in the Transactions of the Royal (Society of 
Victoria, 1892 (4). The hithei'to recorded species are distributed 
as follows: — Ceratella fasca : Coogee, Bondi (N.S.W.), Broughton 
Island, Flinders Island, Lord Plowe Island. C. jiroca.mbeiis : Cape 
of Good Hope, Natal. C. spinosa: Port Natal. Chltiiia ericoj)sis : 
New Zealand. Dehitella atrorahens: Delagoa Bay. In brief, 
the family has hitherto been known to occur only in Australasian 
and S. Afi-ican waters. 

The discovery of a new species of the genus Ceratella in the 
ti'opical waters of the Zanziliar coast is in itself worthy of note, 
but especially so in view of the fact that it is asso(;iated witli 
corals, alcyonarians, and other animals charactei'istic of the 
tropical belt of the East African coast. I am indebted to 
Mr. C. Crosslaud, of Clare College, Cambi-idge, who collected the 
three specimens during his recent exjiedition to Zanzibar, for 
pel-mission to examine and describe them. 

The African species of the family have not yet been accurately 
described, but the accounts of Gray (1) and Carter (2) are 

1 For fX]il!iimtioii of the I'late soc j). 11(3. 

Proc. Zool. Soc— 1903, Vol. I. No. ^^111. 8 

114 PROP. s. J. HicKSON ON [Feb. 3, 

sufficient to show that the Zanzibar specimens should be described 
as belonging to a new species. 

They appear to be most closely related to the Australian 
species Ceratella fusca, but diifer from it in one or two characters 
which Spencer and his predecessors regarded as of generic 

The genus was defined by Spencer as follows :— Colony 
irregularly branching ; more or less expanded in one plane ; 
growing from a creeping base. Main stem flattened, branches 
i-ounded and beset with bracket-like hydi'ophores. 

In C. ifninima from Zanzibar the main stem is not flattened 
but perfectly cylindrical in form, and the hydrophores are so 
extremely reduced or rudimentary that they are little moi-e than 
ridges on the proximal lips of the hydi-opores. The branching, 
moi-eover, appears to be strictly in one plane, and the terminal 
branches are much more slender and delicate than in the other 
species. Before passing on to the specific characters, the size of 
the specimens must be considered. 

The measm^ements are as follows : — ■ 

Specimen A. Specimen B. Specimen C. 
mm. mm. mm. 

Height of the colony 29 22 35 

Maximum expanse of the 

branches 65 38 50 

Diameter of main stem 1"2 0*75 1 

The colonies of Ceratellcc fihsca are from 11 to 5 inches in 
height ; the largest specimens of C . procumbens described by Carter 
were 1 1 inches long by 5 inches broad, and of C. spinosa 4~ inches 
long by 2 broad. The height of Chitina ericopsis is 14 inches, 
with a trunk (main stem) 1 inch in diameter. The size of 
Dehitella is not given by Gray, but from the figures it may be 
judged that it is larger than Ceratella fusca. From these figures 
it is clear that the Zanzibar specimens are much smaller than the 
average size of the adult colonies of the other species. Are they, 
therefore, to be regarded as young colonies or as the representa- 
tives of a dwarf species ? If they are young colonies, it is quite 
possible that the main stem or ti-unk becomes somewhat compressed 
in the plane of branching as the colony grows ; but the fact that 
all the three specimens obtained are of appi'oximately the same 
size, suggests that they have reached or nearly reached their 
maximum growth. The dwarfing of the tropical species of a genus 
that is pi'incipally distributed in temperate waters is not without 
pai'allel in the group of Coelenterata. The very rudimentary 
chai'acter of the hydrophores, however, cannot be explained by 
the suggestion of immaturity, and must be regarded as of specific 
importance. It is true that no gonophores have been discovered 
in the specimens, but it is quite probable that, as in other 
Coelenterates of the tropics, their production is rapid and strictly 
seasonal, so that no ai-gument can be deduced fi'om this character, 
either for oi- against the theory of juvenility. 

The Colony. — The biunching is not very profuse, and strictly 


confined to one plane (PI. XIII. fig. 1). The main stem and the 
larger branches seem to have divided dichotomously with the pre- 
dominance of the most favoiu'ed branch. The terminal branches 
are veiy delicate, branches of 6 mm. in length gradually attenuating 
from 0-2 mm. to 0"1 mm. in diameter. Each terminal branch ends in 
a facultative gl•o^\•ing point, and it appears probable that the growth 
is continuous. I have compai-ed my specimens from Zanzibar 
with a beautifully preserved specimen of Ceratellafusca, for which 
the Manchester Museum is indebted to Pi'of . Spencei- ; and I have 
noticed that the Austi-alian species is much coarser in appearance, 
especially in the region of the terminal branches. I have seen 
nothing in the Zanzibar species corresponding to what Spencer 
calls "The growing ends of the smaller branches" in Ceratella 
fusca, which ai-e flattened in a plane at right angles to that in 
which the general growth takes place and are entirely devoid of 
zooids. It is possible that the diflei-ence may be accounted for 
on the supposition that in Ceratella fasca the growth is seasonal 
or periodic. 

The branches of C. minima are invariably rounded. I have 
seen no evidence of a compression or flattening in any region. 
The sui'face is i-elatively smooth and fi-ee fi'om any spines. Tlio 
hydrophores are represented by very narrow ridges on the 
proximal border of the hydropores. The principal horny fibres, 
I'unning longitudinally with a slightly spii'al twist, may be clearly 
seen through the superficial ectoderm, and in the spirit-specimens 
give a ribbed appearance to the surface (PI. XIII. fig. 2). 

The Zooids are numerous on the terminal bi'anches, less 
numei'ous on the thicker branches, and very scarce on the main 
branches, as in Ceratella fusca. On the terminal branches they ai-e 
ari'anged slightly to one side of the two lateral lines at intervals 
of about one millimetre opposite or alternate to one another. A 
few zooids occur more irregularly distributed. 

On comparing such a terminal branch with one of Ceratella 
fasca, it is at once apparent that in the Australian species the 
zooids ai'e more numei'ous and much more irregulai'ly distributed 
on all sides of the branch. Each fully expanded zooid projects 
about 0'7 mm. from the hydropore, and is about 0*14 mm. in 
diameter. It bears a variable number, but usually nine capitate 
tentacles, each about O'l mm. iia length. 

No gonophores were found on any of the three specimens I 
have examined. The skeleton in the tei'minal branches consists 
of one or two main longitudinal horny I'ods supporting numerous 
looping and irregular bands, which maintain the cylindrical form 
of theVanch (PI. XIII. fig. 3). 

In the larger branches the main longitudinal rods are more 
numerous, and, being chiefly superficial in position, give a slightly 
spiiul, longitudinally striated or ril)l)e<l appearance to the surface, 
in this respect ottering a markeil contrast to Ceratella fasca. 

A series of sections through a small branch shows tliat there is 
present a thin continuous coat of ectoderm covering the whole 
branch, as described and figured bv Spencer in Ceralella fasca, 



The ariungement of the canals and the general histology does not 
appear to differ matei'ially from the desciiption given by the same 
author ; but as the state of preservation of the Zanzibar specimens 
was not perfect, a detailed and critical examination of the sections 
was not made. I was fortunate enough to find, however, a few 
nematocysts that were exploded but I'emained in situ. They are 
(PL XIII. fig. 4) very similar in form to the small nematocysts 
■of Millejjora, and exhibit a vesicle and neck O'Ol mm. in length, 
;armed with four barbs or spines. The thread was invariably 
brolcen, and I have no means of measuring its extreme length. 
The endoderm of the tentacles is solid. 

The diagnosis of the species is as follows : — 

'Ceratella minima, sp. nov. 

Colony probably erect, branching strictly in one plane, iri'egulaily 
and not very profusely. The main stems and all the branches 
cylindrical in form. Hydrophores very slightly developed. 
Skeleton consisting of dark brown longitudinally disposed horny 
fibres, united by loops and bands, forming in the larger branches 
a dense and firm but flexible skeletal plexus. 

Zooids situated slightly to one side of the plane of branching of 
the colony, alternately or in pairs, at distances of about 1 mm. on 
the terminal branches. Lai-gest colony 29 mm. in height, with a 
maximum expanse of 65 mm. 

Locality. Zanzibar, shallow water. 


1. J. E. Gray. — Notes on the Ceratelladte, a Family of Keratose 

Sponges. Proc. Zool. Soc. 1868, p. 575. 

2. H. J. Carter. — Transformation of an entire Shell into 

Chitinous Structure by the Polype Ilydractinia, with short 
descriptions of the Polypidoms of five other species. Ann. 
& Mag. Nat. Hist. ser. 4, 1873, xi. p. 1. 

3. W. M. Bale. — Some new and rare Hydroida in the Australian 

Museum. Proc. Linn. Soc. N. S.Wales, 1889, p. 748. 

4. W. B. Spencer. — On the Structure of Gei^atella fusca. Trans. 

Royal Soc. of Victoria, 1892, p. 8. 


Pig. 1. Drawing of the whole colony of Ceratella minima (specimen A) twice the 
natural size. The base of attachment is covered by an encrusting polyzoou. 

Pig. 2. A portion of a large branch and smaller branchiet taken from the region 
marked * in fig. 1, more highly magnified, showing the polyps P partiallj'- 
extended, bearing an irregular number of knobbed tentacles, t.t. At 7^./^. are 
seen the rudimentary hj'drophores. The figures show the general arrangement 
of the fibres of the horny skeleton. In the large branch they are closely 
crowded togetlier. In the smaller branches the principal fibres are separated 
by considerable spaces but maintain a parallel arrangement. 

Pig. 3. A still smaller branch more highly magnified, in which there is only one 
primary longitudinal fibre. 

Pig. 4. One of the nematocysts of the tentacles. The diameter of the cyst is about 
O'Ol mm. 


8. Contriljutions to our Knowledge o£ the Plankton of the 
Faeroe C-hannel. — No. VIII. ^ By G. Heubert Fowlkii, 
B.A., Ph.D., F.Z.S. 

[Koceivod Dcccmlicr 20, 1902.] 

(Text-figures 13-17.) 

The present pai:)er contains notes {in some cases due to tlio 
vnlued help of friends) on Beroe, Arachnactis, Podon, the Ostra<:'oda., 
the Oopepoda, the Anipliipoda, and the Sehizopoda., captured by 
H.M.S. ' Research' in 1896 and 1897 in the Faeroe Channel. 

Beroe cucumis Fabricius. 

This species, cliai-actei'istic of cold Arctic curi-ents ', was taken in 
the following hauls : — 

16«ii., 300 to 170 fathoms, seven specimens. 

IS e, 400 to ? fathoms, one fragment. 

20 f^, 500 to 400 fathoms, one specimen. 

Most specimens showed the characteristic brick-red or rose tint, 
and though much battered and in some cases inverted, were 
refei'abl'e with a fair amount of certainty to this species. 

A N T II o z A, 
ARACnxACTis ALBiDA M. Sars. 

Some information as to the develojmiental succession of the 
mesenteries in this form was given in ISTo. III. of this series. 
Since its publication, I am glad to say that it has been substanti- 
ally corroborated by Prof, van Beneden ''. 

He agrees with my suggestion to sepa.rate the Channel and 
ISTOrth Sea Arachnactis from alh'ida of the Faeroe Channel, and 
describes it under the name of lloydii, under the idea, Avhich is 
probable, but at present improved, that it will be shown eventu- 
ally to be the larva of Cereanthus lloydii. Till this has been 
proved, I venture to think it better to retain my pro\isional naino 
of hournei for this form. 

The occurrences of A. albida are shown in the table (p. 118) : 
it occurred in over 61 per cent, of epiplnnkton hauls, never in a 
mesoplnnktou haul, and may fairly be taken to be a purely 
epiplankton form. It was present in considerable quantity, as 
many as 50 specimens having been taken in one Jiatd. 

1 The references to previous papers in (lie Society's Proccedinirs are : — No. T., 1800 
p. 991 ; No. II.. 1897, p. o2.3 : No. III., 1S97. p. m:S ; No. IV.,1K98, p.olO; IS',,. V., 
1898, p. 550; No. VI., ISitS, p. 5C.7 ; Xo. VII., 1898, ]>. lOlC. I regret that various 
circumstances, mostly beyond my control, have caused so irreat a lapse ot time 
between this paper and No. VII. 

- Chun : ' Die Ctenoi)horen der Plank ton-Kxjiedition,' p. 2li. 
3 Van Ueneden : ' Anthozoaircs de la Plankton-Expedition.' 



[Feb. 3, 








« r-" 


-2 c 



a 3 




h V- 

S g 



















12 S. 




12 c. 

•)- + 


12 d. 







13 e. 


+ + 


+ + 

13 A. 


13 i. 


+ + 






+ + 


13 Z. 




15 a. 




15 6. 




15 c^. 

16 6. 
16 e. 

18 a. 

19 c. 

19 d. 

20 e. 
20 y. 





+ + 







12 a. 


12 e. 




13 d:. 









16 a i. 










18 6. 



19 a. 





20 a. 





20 6. 
20 c. 





20 i. 







13 a 6. 






13 e. 







15 c. 






19 6. 



















« s5 
S 3 







IS « 






Si Si 



















































































[Feb. 3, 



My friend the Rev. T. R. R. Stebbing, F.R.S., was kind enongh 
to identify and count the specimens of this species. It would no 
doubt have occurred in more of the svirface hauls had not my 
finest tow-net been devoted to the collection of Diatoms for the 
Scottish Fisheiy Board. It is no doubt a purely epiplanktonic 
form in the Faeroe Channel ; a single sjoecimen only was taken at 
20 d (500-400 fathoms) as against about 106 specimens in 7 hauls 
at the svirface ; the single deep specimen was probably a sinking 

This species is, I believe, known only from the surface ^ ; it 
ranges over the Baltic, Norwegian coast up to Vadso, Denmark, 
Boulogne, Concarneau, Trieste. 


The representatives of this order belonged exclusively to the 
Halocypridse, and were mostly taken in the mesoplankton. Only 
one sjDecies, Conchoecia maxima, occui'red sufficiently often to allow 
of a generalisation as to its horizon. 

The identification of a Halocypiid is rarely satisfactory without 
dissection of the mouth-parts, which means destruction of the 
specimen. I have, however, dissected a considerable number, and 
feel at all doubtful only in the case of G. porrecta ; I have pre- 
ferred, however, to leave the specimens under this species leather 
than create a new species on the strength of slight differences 
in the armature of the mandible. 

The table following shows those svirface and deep-water hauls, 
made by Professor Chun between Finistere and the Canary Islands 
with open vertical nets^, which contained the same species as 




Horizon in fathoms. 


1 = 



^ 2 


o ■ 





546 to 




819 to 






546 to 






873 to 



. * 

1 J. de Guerne: Bull. Soc. Zool. Trance, xii. 341 (1887). W. Lilljeborg: 
' Cladocera Sueciaj,' Upsala, 1900, 4to. 

2 Chun : SB. kon. preuss. Akad. Wissensch. (1889 xxx. Claus : ' Die Halocy. 
priden,' Wicn, 1891, 4to. 


were captured by the ' Research ' in the Faeroe Channel ; they 
are cited in the text by Roman numerals. 

CONCHCECIA MAXIMA Brady (fe Norman, 

As I have previously pointed out \ tliis species appears to be 
purely mesoplanktonic, in the latitude at any rate of the Faeroe 
Channel. It occurred in 50 per cent, of the mesophvnkton liauls, 
and in three hauls which began at or over 300 fathoms and finished 
at the surface ; it was not captured once in hauls between 100 
fathoms and the sui-face {cf. table, p. 118). The species was 
fairly common. 

The record of previous captures Avas cited in the second paper of 
this series \ and also indicated a mesoplanktonic habit in subarctic 
regions, but it ls not surprising that in yet colder waters it should 
appear at the surface. At 84° 32' N., 76° E., it was captured 
with a surface-net by the ' Fram'",and is recorded as abundant in 
most of the samples of Crustacea from this voyage ''. 


Twelve specimens in haul 13 i, 100 fathoms to surface. Five 
specimens too small for satisfactory identification, but perhaps 
referable to this species, occurred in hauls 20 c and 20 d. 

Glaus ^ records as other occui-i'ences Chun's haul IV., Ischia at 
492 fathoms, Oi'otava at the sui'face. 


Numerous specimens from 20 a bod and one from 13 i. Claus 
(oj). cit.) records it from Chun's hauls II., III., IV., VII. 

CoxcHCECiA BOREALis G. O. Sars. 

A single specimen in haul 19 «, 480 to 350 fathoms. Recorded 
previously from 250 to 300 fathoms at the Lofoten Islands ', and 
from Trondhjem Fjord at 150 fathoms®. This appears to be a 
purely cold-water form. 

Paracoxch(ecia oblonga Claus. 

Six specimens in haul 20 c, four in 20 (L Claus {op. cit. p. 64) 
cites this species as from Chun's hauls III. and IV. ; and states 
that it also occurs at the surface, but without giving authoiity or 
details. He remarks on the probable identity of this species with 
G. W. Midler's variabilis', a suggestion with which Miiller seems 
to agree "*. This would extend the distrilmtion considerably, as 
Miiller '' records it, from the ' Vettor Pisani ' collections of 

1 Proc. Zool. Soc. 1897, p. 523. 

2 F. Nansen : Norwegian North Polar E.vpuditioii. G. 0. Sai-s: Cnistacua, p. 11. 

3 Id. ibid. p. 137. 

■1 C. Claus, op. cit. p. 61. 

5 G. O. Sars : Fori). Vid.-Selsk. Christiania (1865), vol. 1866, p. 120. 

6 G. S. Brady & A. M. Nonnan : Trans. Koy. Dublin Soc. (2) v. p. 086. 
' G. W. Miiller: " Ueber Halocypriden," Zool. Jahrb. Syst. v. p. 273. 

8 G. W. Miiller : 'Ostracoden des Golfes vou Neapel,' p. 229. 


Chierchia, as occurring at various points in the tropics at depths 
between 382 and 546 fathoms ; it occurs also in the Gulf of 
Naples. Brady ' records it as having been taken by the ' Chal- 
lenger ' Expedition off Kandavu, Fiji, and between Marion and 
Crozet Islands at unrecorded depths. 

Halocypria globosa Claus. 

Six specimens in haul 1 3 i. 

This species is known from the surface and at various depths 
in the Atlantic ^, and is recorded from Gibraltar as taken by the 
' Yettor Pisani ' ^ Of specimens taken by the ' Challenger,' the 
record was in one instance lost * ; other specimens were captured 
at the surface between Api and Cape York \ It seems to be a 
form widely distiibuted both vertically and horizontally. 


Only three complete specimens, and one empty carapace, refer- 
able to this genus were obtained. The specimens on which Claus 
founded the genus (with this single species) were all young males : 
larger specimens of the genus, including females, were obtained by 
Sir John Murray on H.M.S. 'Triton' in 1882, from the Cold 
Area of the Faeroe Channel, and were described by Canon 
Norman and Dr. Brady under the specific name of lacerta, not 
without the " suspicion that they may perhaps belong to the adult 
form of C. daphnoides " ^. My own specimens were too few to 
settle the point ; but as the two smaller specimens most resembled 
in outline the figvire of Claus, and the two largest that of Brady 
and Norman, I have left them provisionally under the older specific 

In addition to Chun's hauls III, and lY., it has been captured 
at 200 fathoms off Achill Head {daphnoides) ^ the Faeroe Channel 
as above {lacerta), and off Kandavu, Fiji, at an unrecorded depth ^ 


Mr. I. C. Thompson was kind enough to report on the 
Copepoda in No. lY. of this series of papers '^. Since that date, 
the arrangement of three then doubtful hauls has required modi- 
fication : 12 a, which was suspected at the time of capture to have 
remained open too long, proves to have no apparent contamination 
of undoubted surface forms, and has been moved to the Meso- 
plankton ; 1 3 e was also suspected, in this case with justice, as 

1 G. S. Brad}' : " Myodocopa of the ' Challenger ' Expedition," Trans. Zool. Soc. 
xiv. p. 95. 

2 C. Claus, op. cit. p. 79. 

3 G. W. Miiller : Zool. Jahrb. Sysfc. v. p. 270. 

* G. S. Brady & A. M. Norman : Trans. Roy. Dublin Soc. (2) v. p. 705. 

5 G. S. Brady : Trans. Zool. Soc. xiv. p. 97. 

6 G. S. Brady & A. M. Norman : Trans. Roy. Dublin Soc. (2) v. p. 697. 

7 G. S. Brady : Trans. Zool. Soc. xiv. p. 95. 

8 Proc. Zool. Soc. 1898, p. 540. 


containing several undoubtedly epiplanktonic species (e. g. Arach- 
nactis albida), and has been relegated to the " doubtful" category, 
closure of the net not having taken place at the proper time ; 
12/ was known to be epiplanktonic, although there was some 
doubt as to the exact depth at which it had been towed. I have 
therefore reprinted in the table (pp. 118, 119) the captures of the 
seven fonns which were taken at least six times, sufficiently often 
to give ajDproximate data for an estimate of their vertical distri- 
bution. In discussing this question, I gave a short table on 
p. 546 showing the occurrences of these seven species expressed 
in percentages of those Epiplankton and Mesoplankton hauls 
which contained Gopepoda ; this can now be amendetl as follows, 
omitting the four doubtful hauls from the calculation : — - 

Total hauls containing Copepoda. 





Calamisfinmarchicus occurr 

ed in 

83 "/n 



91 "/o 

^ Eucalanns attennatas 

22 „ 

41 „ 

lEiicJueta norvegica 

11 » 

75 „ 

3[ctridia longa 


75 „ 

'^ Pleuromma ahdomiiiale 

"5 ,',' 

58 „ 

Acartia clausii 

33 „ 

25 „ 

Temora longicornis 

33 „ 


The amended table is in harmony with the conclusions dnxwn 
from the former, as to the vertical distribution of these forms, 
except in the case of Acartia clausii, the question of which was 
expressly resei-ved {pp. cif. p. 549). 

Since the publication of Mr. Thompson's report, I found that 
Dr. R. Norris Wolfenden was making an exhaustive study of the 
fauna of the Faeroe Channel, and natumlly j^laced my collection 
at his disposal. He has been kind enough to f ui^nish the following 
notes on new and other species, with some of which he has 
already dealt briefly elsewhere ^, Exact data of depth &c. were 
not always available, as by the time that Dr. Wolfenden received 
the specimens all the epiplankton hauls of a station had in many 
cases been put together in one bottle, all the mesoplankton hauls 
in another, for economy of space. 

Pleuromma robustum Dahl, Zool. Anzeig. v. p. 16 (1893). 

"This is the common Fleioromma of the Faeroe Channel, and 
was found to be present in considerable numbers in Dr. Fowler's 
collection, PL ahdominale occurring much more rarely. In these 
northern latitudes it nlmost entirely replaces the latter species. 
It is readily distingviishable by the horseshoe-shaped mass of red 
pigment which is present in the anterior and inferior portion of 
the head at the base of the mouth-organs, and in the male by 

^ Dr. Wolfenden points out that as, according to his wide experience, T^ucalanus 
attenuatus is nowhere met with in the Faeroe Channel, unless perhaps quite 
exceptionally, these figures probably refer to Eucalantis elonijatiis. Similarly, for 
the species Pleuromma ahdominale should probably be substituted P/. robustum. 

- R. N. Wolfenden : Journ. Marine Biol. Assoc. (1902), vi. p. 344. 


the clasping antenna being on the left side, and the pigment- 
spot invariably on the right side. The secend feet in both 
male and female have the characteristic notch and hook on both 
limbs. Length 3-4 mm. Haul 20 a (200 to 100 fms.) and several 

HetergcHtEta zetesios c5" , Wolfenden, op. cit. p. 367. 

" The head is like E. jyapilUgera Gbt. Though the end joints 
of both anterior antennse were broken off, the 19 joints left had 
a length of over 4 mm., with the geniculation between the 18th 
and 19th segments. The anterior antennae were therefore much 
longer than the whole animal, which was 3*5 mm. There was 
considerable asymmetry of the furcal segments, that on the left 
being much the longest and broadest. The anterior foot-jaw had 
one thick hooked bristle on the 5th lobe, but no " tooth-comb " 
bristle, and the 5th feet were peculiar and unlike those of any 
other Heterochceta, displaying an upright and stiff process of the 
2nd basal joint, armed with fine stiff hairs on the inner aspect 
(like a " tooth-comb "), and the proximal inner margin of the 1st 
joint of the exopodite with a protuberance armed with 4 teeth. 
The 2nd basal joint of the foot of the opposite side is armed distally 
with short stiff bristles. It could be only the male of H. grimaldii 
or of H. longicornis, neither of which is yet known, or of H. major 
(Dahl). The latter and R. grimaldii are very large (5-10 mm.), 
and though II. zetesios resembles II. longicornis in some points, it 
is perhaps better for the present to distinguish it as a new species. 
Only one example was met with in Dr. Fowler's collection, in 
haul 20 « (200 to 100 fathoms)." 

^GiSTHUS ATLANTicus Wolfenden, op. cit. p. 364. 

" The occurrence of an example of this genus in the Faeroe 
Channel is remarkable. This specimen was found in the collection 
made by Dr. Fowler as 20 a (200 to 100 fathoms). It had a 
total length of 1*45 mm., a 6-segmented anterior antenna with 
very long and peculiar sensory processes. It has distinct differ- 
ences from uEg. 'mucronatus or ^Eg. acideatv.s Gbt., and also from 
the species described by Scott from the Gulf of Guinea as 
jEgisthus longirostris." 

LucicuTiA MAGNA Wolfenden, sp. n. c? • 

"A single specimen found in Dr. Fowler's collection from 19 a 
(480 to 350 fathoms), of 3*54 mm. length, was apparently new. 
The anterior antennas were larger than the whole body, by the 
terminal one and a half joints. The endopodite of the 1st foot 
was two- jointed. The right 5th foot has a strong spiny process 
on the inner side of the 2nd basal and the exopodite of two seg- 
ments ; the endopodite and exopodite of the left 5th foot being 
■ each of three segments. The size alone distinguishes it from the 
males of any other known species, only L. grandis being larger." 


AuGAPTiLus ZETESios Wolfeuden, op. cit. p. 369, 

"One specimen only was found, in the bottle marked 19 a. c. 
Another sijecimen occurred in the sample marked 20." 

EucALANus CRASsus Gbt. 1888, Atti Ace. Line., and 1892, Fauna 
u. Flora ISTeapel, v. p. 1 9. 

" This species was of not infrequent occurrence, especially in 
the bottles marked B 1 and 13 A- (2 to 0) and 13 i (100 to 0). 

" The writer also has frequently noted its occurrence in the 
Faeroe Channel." 

Gaetaxus major Wolfenden, sp. n. 

" Two examples of this genus {Gaetanus) were found in 
Dr. Fowler's collection marked 19 « (480 to 350 fathoms). The 
copepod very greatly resembled Gaetaims armiger Gbt., but the 
anterior autennc-e were longer, i-eaching beyond the f urea by the 
length of the last joint ; the spines of the last thoracic segment 
were comparatively shorter, the 1st abdominal segment and tlie 
anal segment shorter, and the furcal segments only as Ion"- as 
broad (longer than broad in G. armiger), and each abdominal 
segment had a row of pectinations on the posterior border. The 
saws of the swimming-feet possessed more teeth ; the abdomen 
Avas not nearly half the length of the cephalothorax, and the 
whole length of the animal was 5-3 mm. In all these points it 
difiered from the typical G. armiger, the size of which reaches 
only about 3 mm., and justifies its being made into a sejjarate 

Gaidius Gbt. 1895, Bull. Mus. Harvard. 

" A good many examples of this genus occurred in the deep- 
water collections of Dr. Fowler, e. g. in the bottle marked Meso- 
plankton 20 (500 to 100 fathoms). 

_ There is no doubt that in the Faeroe Channel there are two 
kinds of Gaidius — one agreeing in every particular with the 
Gaidias 2iangens oi Giesbrecht; the other, a larger s^oecies, also 
difiering in the segmentation of the 1st and 2nd feet. Gaidius 
2)iingeas Gbt. has the exopodite of the 1st foot with only two 
segments ;ind the endopodite of the 2nd foot with only one seo-- 
ment; whereas the northern species has a 3-jointed exopodite 
of the 1st foot and a two-jointed endopodite of the 2nd foot. 
There are other minor differences. In size the northern species is 
much larger ; Gaidias jningens Gbt. being 4| mm,, tis compared 
with about 3 mm. The Chiridius temcis2nnis~'oi G. 0. Sars is the 
same species, all being characterized by the peculiar series of 
lamellar ajipendages of the basipodite of the 4th foot." 


" Some examples of this species occurred in Dr. Fowler's col- 
lection. It was drawn and described (in IM8S. only) before the 


writer became acquainted with the recent work of Prof. G. 0. Sars, 
who figured and described the species as Chiridius armatus. The 
writer has pubhshed reasons why this generic name should not 
be used (see Rep. of the Brit. Assoc. 1892, " A proposed Revision 
of the Subfamily Aetidiince "), as it is not a Chiridius." 

EuCHiRELLA CARINATA Wolfenden, op. cit. p. 366. 

" One example of this species, measuring 3*54 mm. in length, 
was found in Dr. Fowler's collection in the bottle marked 20. 

"These new species will be described in full, along with the 
drawings, in the writer's monograph which is in hand." 


I am indebted to the Rev. T. R. R. Stabbing for help in the 
determination of some of these forms. Only one species occurred 
in sufhcient quantity and with sufficient frequency to enable 
deductions as to its habitat in the Faeroe Channel being drawn, 

Parathemisto oblivia 'Krojer = abi/ssorum Boeck. 

I have already discussed the distribution of this form at some 
length ^ ; to the records there given must be added two stations 
of the ' National ' expedition — N. of the Hebrides, and S.W. of 
Iceland '^ — and 1 2 stations along the route of the ' Fram ' ^ Canon 
Norman ajso cites Bonnier as having taken it at 950 meti-es in 
the Bay of Biscay *. 

It is apparent from the table (p. 119) that the species is a 
true member of the Mesoplankton in this locality, having been 
captured in 66 per cent, of the deep hauls ; it rises to the surface 
at midnight, the only occasion out of 26 Epiplankton hauls being 
at that hour (haul 1 5 d). 

Cyclocaris guilelmi Chevreux. 

Two specimens of a Cyclocaris were obtained in haul 20 d, 
between 500 and 400 fathoms. Mr. Stebbing informs me that 
they agi'ee undoubtedly with the above species, captured in a net 
sunk to 600 fathoms near the Lofoten Islands by the ' Princesse 
Alice ' '% the largest of the six specimens being about 1 2 mm. in 
length. My largest specimen, a female, would have measured 
about 20 mm. if straight. 

Another species of this genus, Cyclocaris tahitensis Stebbing, 
was described ^ from a single specimen taken by the ' Challenger ' 
off Tahiti, apparently at the suiface ''. 

1 Proc. Zool. Soc. 1898, pp. 583-585. 

2 J. Vosseler : Amphipodeii der Plankton-Expedition, p. 80. 

^ P. Nansen : Nonvegian Nortli Polar Expedition. Crustacea, b}' G. 0. Sars, 
p. 14. 

4 A. M. Norman : Ann. Mag. N. H. (7) v. p. 131. 

5 E. Clievreux : Bull. Soc. Zool. France, xxiv. (1899), p. 148. 

* T. R. R. Stebbing : Chall. Rep. Zool., Amphipoda, p. 661, pi. xvii. 
^ J. Murray : Chall. Rep., Summary of Results, p. 1077. 


Mr Stel)biug points out "that tliis species has been a-ain 
figured in great detail hy Pi-ofe,ssor Sars \ and tluit Dr. Nornmn ■= 

T \T^ if , !°'* f'^"'"' ^''''''''' I't^I'i'^s^^ted his Ci/clocarisfaroensis. 
in Mr. btebbings opinion the diflerences between the two sets of 
hgures are purely casual, depending ou individual or accidental 
conditions of the specimens examined. Further, wliile acceptino- 
provisionally the distinction between the boreal form and the 
Cliallenger C. tahitensis, he agrees with the view thus exin-essed 
by i)r. Norman, 'so remarkable is the resemblance, that the 
diflerences seem scarcely varietal ; but I hesitate to unite a form 
Sliti"" Channel with one from so distant a locality as 

Canon Norman does not appear to have known of M. Chevreux's 
name (which apparently has priority); his specimens were captured 
tTa i'!i '^^'V. "' *^'® ^'^^^'^^ Channel in 1882, at a of 
640 fathoms Professor Sars's specimens were apparently from 
various depths along the course of the ' Fram,' the smallest 
specimens at the le-A&t depths. 

This species (omitting tahitensis as specifically distinct) is 
evidently a Polar and deep-water form. 


A single specimen from haul 13 i, 100 fathoms to the surface, 
was Identified by Mr. Stebbing. The previous records of its 
occurrence are: Hardangerfjord, 100 fathoms'; from Ninsen- 
jord to Hardangerfjord^; Folgero, Sunde, Foldenfjord, 80 to 
100 fathoms fears ' regards T. nordeuskioldi Boeck and T. boecki 
fetebbmg as being the males of T. malmi. Of these the former * 
was described from the Sofia Experlition as oflHhe Faeroe Islands 
at 65 N., the latter ^ from 18° 8' N., 30° 5' W., at the surface. 

EuTHEMiSTO sp. indet. 

A number of specimens too much broken for recognition 
occurred m the haul 1 5 c^ at midnight. eco^nition 

EuTHEMiSTo BisnxosA Boeck. 

Occurred in 15 c (530 to fathoms) and Vo d (surface, at mid- 
night). The records of this species were exclusively Arctic, till 

tJZ '^'^9 «T .1^ the 'National' expedition in the Sargasso Sea 
between 218 fathoms and the surface.' 


A single specimen at 12 6 (450 to 320 fathoms). This also was 
believed to be exclusively an Arctic form, but has now been 

pis. H: & "f" '- ^'''''^'-^'' ^°^'l' l'«^-' Expedition. G. O. 8ar. : C.-ustacca, p. 20, 

3 i- ^k ^"vr""/- "^""i ^'*- ^- "• (' ) '-■ V- 197, 1.1. vi. 

, "°«,f -■ J'kaiidiiiaviske og Arktiske Ampliipoda,' 1872, p 92 

;« C. Hovalhus: Vega-Exped. Veteiisk. laktta- iv. p 573 ' 

•' hurs: Cnistafi'M of Norway:, i;p. 17 (189U) 

" T. K. II. Stebbing: Chull. Itop. AiuphipoJa, p. 1539 


I'ecoi'ded from the Antarctic region, as well as from the Gulf- 
Stream pi-oper, the Sargasso >Sea, and South Equatorial Drift ^. 


A single specimen in haul 15 cZ at the surface at midnight. It 
is widely distributed over the Arctic Seas, but, unlike the two 
foregoing species, was not taken in southern waters by the 
' National.' 


A single specimen from 13 g (465 to 335 fathoms) was acci- 
dentally included among the Gopepoda sent to Mi\ Thompson 
and identified by Mr. A. O.Walker. According to Ganon Norman ", 
the disti'ibution of this species is mesoplanktonic, mostly northern, 
but also in the Bay of Biscay (960 metres). It reaches to about 
80° N.' 


The Schizopoda captured belong exclusively to the Euphausiacea, 
and are referable to only three species. Several forms which 
might have been reasonably expected among the captures were 
absent. A list of the British species with their distribution is 
given by Ganon Noiinan, in his paper on Bi-itish Lophogastridse 
and Euphausiidse *. 

Thysanoessa loxgicaudata Kroyer. 

A considerable number of specimens of this species were 
captured : it appeared to be the commonest Schizopod of the 
Faeroe Ghannel at the time. The synonymy appears to be 
Tliysanoi^oda longicaiidata Kroyer = Thysanoessa tenera Sars = 
Thysanoessa longicaudata of Hansen, Norman, Ortmann, &c. 

The species ranges from the West Goast of Norway right across 
to Greenland and into the Labi-ador current ; the ' National ' " ceased 
to take it (after almost daily captures up to that moment) from 
the date of entering the warm water of the Gulf-Stream (' Florida- 
strom '). Sars **, who described the species fi'om deep water in 
the Varanger Fjord, records it also as from the surface at four 
stations between Norway and Jan Mayen on the cruise of the 
' Voringen ' '^. It appears, therefore, to be essentially a cold- 
water species, an Arctic type-form, and was captured by the 
' Fram ' \ 

It has, however, been recorded twice from British coasts °. 

^ Vosseler : Amphipoden der Plankton-Expedition, p. 86. 

2 A. M. Norman : Ann. Mag. N. H. (7) v. p. 135. 

^ F. Nansen, op. cit. p. 19. 

■* A. M. Norman : Ann. Mag. Nat. Hist. (6) ix. p. 454. 

5 A. Ortmann : ' Decapodcn und Scliizopoden der Plankton-Expedition,' p. 14. 

6 G. 0. Sars: Porliandlinger Videnskabs-Selskahet (Christiania), 1882, no. 18, p. 53. 
' G. 0. Sars : Norwegian North Atlantic Expedition (Crustacea), pt. ii. p. 13. 

^ ¥. Nansen : Norwegian North Polar Expedition. G. O. Sars : Crustacea, p. 14. 
" A. M. Norman : Ann. Mag. Nat. Hist. (6) ix. p. 463 ; and the papers there cited. 


Once it occuiTcd in cnoi'inous quantity in St. Andrew's JJay in 
company with Nyctiphanes norvegica, once at Rodcriii- witl> a 
similar swarm of E athemisto comj)ressa, the lattei- l)eing also an 
Arctic type-form '. In both these cases it is probable that the 
creatures had been driven down the North Sea by a .strong southerly 
current, in the manner which I have already suggested '" foi* 
Parathemisto ohlivia; and it has thei-efore no more i-ight to lie 
i-egai-ded as a " British " species than an occasional Velella ov 
lanthina bi'ought up by the North Atlantic Drift to our shoi'es. 
According to Ortmann {pp. cit.), the ' National ' haids gave no 
indication of the vertical distribution of this species. 

It will appear from the ta))lo (p. 119) that the ' Research ' was 
more foi'tunate, and the haids point to its having a distinct 
pi'efei'ence for the mesoplankton in the Faeroe Channel. Like 
(at any lute some) other mesoplaidvtonic species, it rises to the 
surface at night. Specimens with adidt chaiucters were captiired 
in 19 per cent, of epiplankton hauls, but in 06 per cent, of meso- 
plankton haxds. On the other hand, lai'val and j'ost-lai'val stages, 
apparently refeiulile to this species, wei'C obtained in 38 per cent. 
of epiplankton hauls, but oidy in one mesoplankton haul, and 
that one terminating near the 100 fathoms. The species, there- 
fore, appears to be epiplanktonic when young, mesoplankton ic 
when adult, so far as these observations go and in the Faeroe 
Channel at this time of year. In seeking deepei" (coldei-) water 
in this locality, it follows what appeal's to be the practice of other 
Arctic type-forms when they meet the warmer water of the North 
Atlantic J)i-ift. That this w^as not appai-ent from the results of 
the ' National ' is probably due to the fact that from the Hebi'ides 
almost up to the moment of coming into the Gulf-Stream the 
vessel w'as in far colder surface-water than that of the Faeroe 
Channel in summer. 

The larva2 mentioned abo\"e ranged fiom an early Cahjptopis 
stage up to the adult condition. It was not, of course, possible to 
derive them all with cei'tainty from Thysaiioessa longicaudata ; but 
the majority may l)e safely I'efei'red to this species, not only 
because the adults captured were far in excess of any other 
Euphausid, but also because the larva; could be traced gradually 
thi'ough successive stages back to the Cahjptopis. The meta- 
morphoses of this species follow the lines indicated by Sars ^ for 
Nyctiphanes, Eujyhausia, and Thysanopoda. 

As Paul Mayer' has shown, the spination of the telson of 
Malacostracan la.rva; yields a character important both for ph5do- 
geny and for diagnosis. It has not as yet, I think, been pointed 
out that the condition of the telson in Euphausiida? affords a 
further argument for the view maintained by Pxias' and othei'S, 

' C. Chun : ' ]?czichui)gcn '/.wisclicn dcni avktisclicn nml aiit;ukfisilirn I'lnnktciii." 
Stuttgart, 1897, 8vo, p. 30. 
- Proc. Zool. Soc. 1898, p. 583. 
3 G. 0. Sars : Cliall. K.'p. Zool. xiii. (Schizopoda). 
•• P. Mayer: Jpnaischc Zcitsclirift, xi. (1877), p. 210 ct seqq. 
^ .J. E. V. Boas: Morpliologisches Jahrbucli, viii. p. tSo. 

Proc. Zool. Soc— 1903, Vol. I. No. IX. 9 


that this family takes its origin very near to the root of the 
Decapodan stem, and that it has far closer affinities with the 
latter than with the Mysidse. In Nyctiphanes ^ and Euphausia ^, 
and possibly in other genera also, the youngest Calyptopis-laxvse 
show seven spines on each side of the telson ; unlike all other 
Schizopoda, so far as is known, except perhaps Lophogaster, they 
are thus in absolute accord with the " urspriingliche Borstenzahl 
7 + 7 " which Mayer attributes to the primitive Macruran and 
Brachyuran. In Euphausia, Nyctiphanes, Thysanopoda, and 
JVematoscelis according to Sars (op. cit.), and in Thysanoessa, 
the number is increased at later stages by a median terminal 
spine, which, like the others, is jointed to the telson. Accepting 
Mayer's enumeration of the spines from the middle line outwards, 
and styling the median azygos spine of the Euphausiidse as 0, — 
spines 7 are found in the adult Thysanoessa about one-third of 
the length of the telson from the root ; spines 6 at about two- 
thirds of its length from the root ; spines 5 are lost ; spines 4 
persist as the large lateral jointed spines near the end of the adult 
telson^; and spines 3, 2, 1, disappear altogether in the course 
of development. On page 131, I have illustrated four stages in 
this reduction omitted by Sars, of which fig. 15 does not quite 
bear out his description : these show the disappearance of the 
median spine 0, and the commencement of a new unjointed 
growth of the telson backwards, to form the lanceolate tip of the 
adult. The character of the telson and the presence of this 
median spine will apparently form a good criterion for the 
separation of Euphausidan larvae (at stages later than the Meta- 
nauplius) from other Schizopodan and from Decapodan larvae. 

The earliest Calyptopis-lsirvse captured by the ' Research ' 
resembled closely those figured by Sars {op. cit.) for other genera, 
except for the facts that the carapace was much more globular 
anteriorly and was devoid of spines or processes. 

Nyctiphanes norvegica M. Sars. 

This form was captured on only six occasions. Although a 
North Atlantic type, it is not an essentially Arctic type like 
Thysanoessa longicaxidata : it is of constant occui'rence in certain 
localities on our own coasts, and has been recorded from as far 
south as Portugal. The various records of its occurrence are cited 
by Canon Norman *, but unfortunately the size of the individuals 
and the depth from which they were derived are only rarely noted. 
I am informed by Sir John Murray that, in his experience, large 
adult specimens are taken only in deep water. 

1 G. 0. Sars : Chall. Eep. Zool., xiii. Schizopoda, pi. xxvii. fig. 6. 

2 C. Glaus : Untersucli. Crustaceen-Systems, pi. i. fig. 2, Wien, 1876, 4to. 

3 With regard to these, Boas {oj). cit. p. 523, note 5) has suggested that they may 
he homologous with the long caudal appendages of Nehalia and many Phyllopods. 
This possibility is rendered considerably more remote by their being merely two 
persistent spines out of a series which is not represented in the forms cited by him. 

4 A. M. Norman : Ann. Mag. Nat. Hist. (6) ix. pp. 459^60 (1892). 




Text-fig. 13. 

Text-fio- 15. 

Text-fiff. 16. 

Text-fio-. 17, 

Telsoiis of l;ii'v:i' attribiitcd to Thiiaavocxsa Innf/icandald, sliowiiii;' vi'ductioii of tlic 
primitive seven imirs of s|)iiies and torniatioii of tlie median uiijointed sjiine 
of tlie telson. 

Fig. 13. Early Calijptopis, 1'5 nini. long. 

14. JPurcilia, 4"5 unn. long. 

15. Ci/rfopia, 5 nun. long. 

Ui. Late ('uvlopia, 5 mm. long. 
17. 4 mm. long. 


The same distribution was noted by Vallentin and Cunning- 
ham ^ : — " The adult, so far as our information allows of a 
decision, lives on the bottom, and never swims far from the 
ground [90-95 fathoms in this instance], while the young, up to 
half or three-quarters the size of the adult, occur abundantly at 
the very surface and at all intermediate depths. As mentioned 
above, Mr. Murray found swarms of individuals at the surface in 
the Faroe Channel, but none of these were full-grown, and very 
few more than half the adult size." 

I have no doubt that this generalization will prove true for 
greater depths : I took adult specimens, over 35 mm. in length, 
only between 350-220 fathoms, 400-300 fathoms, 500-400 
fathoms: the remaining specimens varied from 9 to 17 mm. in 
length. In other words, Nyctiplianes norvegica is apparently 
mesoplanktonic when adult. 

A few larvse, larger for their stage of development than those 
attributed to Thysanoessa, were taken at the surface, and may 
perhaps belong to JSfyctiphanes : they have not been included in 
the tables. 

Thysanopoda microphthalma Ortmann. (? = Thysanopoda 
mia'ojihthaVma G. 0. Sars.) , 

Three specimens, recognizable as young forms by the character 
of the second maxiUa and gills, and by the spination of the telson, 
of about eight, twelve, and fourteen mm. in length, appear to be 
referable to the same species as specimens recorded in quantity 
by the ' National ' in 60° 3' N., 27° 0' W., at a probable depth of 
between 218 and 328 fathoms. These were referred by Ortmann" 
to Thysanopoda 'microphthahna of Sai'S ^, a species founded on two 
specimens from the surface at 26° 21' N"., 33° 37' W., and 7° N., 
23° W. respectively. The identity of the ' National ' specimens 
with those described by Sai'S seems to me rathei- doubtful : firstly, 
because it is not very likely that a rare form svich as this should 
occur as adult both at the surface near the Equator and also at 
218-328 fathoms in the Greenland Sea, or at 500-400 fathoms 
in the yet colder water of the Faeroe Channel ; secondly, because 
Ortmann himself indicates some points of difierence between his 
specimens and those of Sars. My own specimens agree with 
Ortmann's figure, and dififer from Sars's description, in the shape 
of the antennal scale, and in the absence of a spine from the 
second joint of the first antenna. The telson was not hispid, 
probably owing to immaturity ; the eye was somewhat flatter than 
in Ortmann's figure, and showed slight signs of a constriction 
such as is charactei'istic of Thysanoessa. The matter cannot be 
settled in default of further specimens, owing to the fact that 
Sars gave only a woodcut of the entire animal, and no figures of 
the detailed anatomy. 

1 R. Vallentin & J. T. Cunniugliam : Quart. Journ. Micr. Sci. xxviii. pp. 325-6. 
- A. Ortmann : ' Decapoden und Schizopoden der Plankton-Expedition,' p. 9. 
3 G. 0. Sars : Chall. Rep. Zool., xiii. Schizopoda, p. 106. 

1903.] ON THE ELK IN NORWAY. 133 

Of the tliree ' Research ' specimens of this species, two came 
from the Mesoplaiikton, one from a haul of 480 to fathoms : 
no examples were captured at the surface, unless some of the 
larvae attril)utefl to 7"/^ ?/,s-rtv(oessa belonged to this species ; this is 
imlikcly, liecause in tliat case the small size of the eyes would 
prol);il)ly have l)etrayed them. 

[). On the Present Condition and Habits o£ the Elk in 
Norway. By H. J. Elwes, F.R.S. 

fKeceivcd January 19, 1903.] 

(Text-figures 18-26.) 

tSo little seems to be known l)y naturalists concei'ning the actual 
condition of this remarkable animal, that I think some of the 
observations I have made dui-ing six Septembeis, which I have 
spent entirely in the fascinating spoit of Elk-hunting, may be 
worth recording in the Proceedings of this Society. 

My experience has been gained entii-ely in the pro^'inces of 
North and South Ti'ondhjem, where the Elk is more numerous 
than pci'haps in any other pait of Europe, and where it seems to 
attain a. gi'eater size and vigour, if one may judge liy the develop- 
ment of the horns (see text-fig. 18, p. 134) than anywhere else in 

I have also derived much vahial)le infoi'mation fi'oui Capt. Geiurd 
Ferrand, who was, I think, the first Englishman to liunt regukxrly 
in the same districts, and whose experience continued ahuost 
witliout a break for 20 years fi'om 1865. This has been invaluable 
in confii-ming, and to some extent modifying, my own observations 
and what I have been able to learn from the natives of these 

Foi'ty years ago the Elk was a much scarcer animal in Ceutiul 
and Northern Norway than it is now, or i-athei" was ten years ago, 
since whicli time it has probaljly decreased in numbers. At that 
time it was hunted for its meat alone by the proprietors of tlie 
regions where it occurred, and in the lai-ge tracts of forest 
belonging to the Government it was not allowed to be shot until 
1880. Up till that time very few foreigners knew what magni- 
ficent sport Elk-hunting was, and as the natives preferred the 
meat of cows to that of bulls, which, after the rutting-season 
begins, is extremely rank and hard, the bulls increased in numbers 
and were able to grow to a size and age which they rarely 
attain now. 

The legal season for killing Elk was formerly much longer 
than it is now, and in North Trondhjem lasted for three months. 
Then it was cut down to 6 weeks, from the 1st September till the 
15th October, and now has been still further reduced to a, month 
or 3 weeks in some j^rovinces, Avhilst in Sweden only 15 days are 
allowed during which the animals can legally bo killed. 



[Feb. 3, 

2 03 

i2i n 

Jzi p 

'5 2 




Text-fiff. 19. 


Horns of fully adult Elk killed in Bjoindal, 65° N., Sept. 4 1895 ; 18 points; a fail- 
average size for this district, but uuich larger than usuallj' seen in the south, 
(y'l nat. size.) 

Text-fig. 20. 


Horns of a full-grown but not adult Elk, killed Sept. 1896 in Upper Namdalen, 65° N. 
Supposed by natives to be 4 or 5 years old. Probably this type would develop 
into such antlers as shown in text-tig. 18 if conditions were favourable, 
(ji nat. size.) 

Text-fig. 21. 

of young Elk, sui)]>oscd to be 3 years old. These would probably develop 
into such a head as that shown in text-fig. 19. (yf uat. size.) 

136 MR. H. J. ELWES ON [Feb. 3, 

This rapid increase of an animal which, on account of its great 
size and conspicuous tracks, cannot escape unnoticed in an 
inhabited country, must be entu-ely attributed to the wise game- 
laws made by the Norwegian Government, and in most parts 
honestly observed by the most law-abiding and well-governed 
people I have ever met with. In former times the Elk, in Scandi- 
navia, as in ISTorth America, was hunted down in winter on ski 
(the Noi'wegian form of snowshoe), and slaughtered for its meat 
by every peasant farmer, till it had almost been exterminated. 
When, however, a law was made that it could be hunted only in 
the month of September, which period was for a time somewhat 
extended in North Trondhjem, its numbers soon increased, and 
about 20 years ago attained such proportions that English and 
German sportsmen began to visit Norway to hunt Elk. The 
landownei's in some districts then discovered that, instead of 
hunting themselves oi- paying Swedish hunters, who, from long 
experience, were more expert, to kill their Elk for them, the 
right of killing Elk, which is limited to one animal on each farm, 
had a letting value ; and when many such rights are united so 
that a large tract of country can be reserved to the lessee, this 
value was worth a little trouble to maintain. The consequence 
has been that, though poaching and killing Elk out of season is 
not entirely unknown, yet it cannot be carried on extensively ; and 
I have little doubt that the Elk will continue to thrive wherever 
the country is suitable. 

The statistics which I append show the numbei's of bull and cow 
Elk which are known to have been legally killed in the various 
" Amts " or provinces of Norway in 1894, and the average for the 
previous five years, and may be taken as very nearly exact, though, 
according to some, the returns for the southern provinces a,i-e not 
so accurate as for the northern ones, and this record takes no 
account of those illegally killed. Tt will be seen that only four 
provinces of Norway (excluding Finmark), namely, Stavanger, 
North and South Bergenhus, and Romsdal, all of which are on 
the south-west coast and exposed to the warm and wet influence of 
the Gulf-Stream, are without Elk ; and it may be added that in 
those four provinces wild Reindeer are most numerous. 

North Trondhjem is before all the rest in numbers ; and in this 
province I believe the size of the horns is or was also much larger 
on the average than in any of the southern districts (text- fig. 18, 
p. 134). 

It is said that the Elk is gradually extending its range north- 
wards, and has appeai-ed in the southern parts of the province of 
Nordland only in the last few years ; and there seems to be no 
i-eason why it should not go still further, as in North America the 
Moose (which is so nearly allied to the Elk, that I do not think 
it can be looked on as more than a variety of that animal) is 
found in regions where the climate and food is certainly not more 
favourable to its habits than they seem to be in some parts of 
Nordland and Finmarken. 

The greater part of North Trondhjem and a large proportion of 

1903.] rut; klk ix norway. 137 

South Trondhjem are mountainous, barreu, and thinly populated 
except on the coast and in some inland valley.s and fiords, and 
are covei'ed with forests of birch, spruce, and Scotch la- up to an 
elevation of about 2000 feet, wherever the ground is not too rocky 
or swampy for these ti'ees to grow. Above that elevation there 
are fjelds or bare mountainous ujjlands, the lower slopes and 
sheltered dells in which are more or less clothed with birch, willow, 
mountain-ash, and alder, mixed with stunted firs. There are 
many large lakes and lai-ge are;is of peat-l)og ; but these bogs are 
rarely so deep anil soft as to be impassable, and even after long 
periods of rain a, man can cross them by picking his way. 

The country is divided into so-called farms, most of which con- 
sist of small patches of oats and potatoes, with from 10 to 30 acres 
of mea,do\v-land which is mown for hay, and the produce of which 
is eked out by small ricks which are put up wherever a, sledge- 
load of grass can l)e got together in the forest. A lai-ge area of 
forest and mountain is attached to ea,ch farm, and there are usually 
one or two sietei-s ^ on the mountain, to which the cows are driven 
for about three months in sunnner. Though most of these farms 
belong to the occupiers there are some large private properties, 
belonging to timber companies and public institutions, and nnich 
of the higher fjeld and barren mountain remains in the hands of 
the Government. 

In the wilder and less populated districts the Elk ai'e hai'dly 
disturbed during eleven months of the year except by the occ;isional 
attacks of bears and wolves. Though the bear has l)ecome rarei' 
of late years, the wolf, on the contrary, has appeared in distiicts 
where it was until recently almost unknown ; but at present 
they have confined their attacks rather to the semi-wild rein- 
deer, which are kept in some numbers by the Lapps all along 
the Swedish frontier, fi'om Roros northwai'd, and to the sheep, of 
whicli every farmer has from 10 to 30. It seems to be doubtful 
whether bears can kill full-grown Inill Elk", lait during my last 
hunting-season I found the remains of no less than three Elk calves 
which had been killed by them. 

The favourite food of the Elk in summer consists almost entirely 
of the twigs and leaves of birch, willow, and mountain-ash, and 
in winter of the branches and bai-k of the same shrubs. Wherevei- 
there is a grove of mountain-ash, Elk will live almost entirely on 
it so long as they can get it, and in disti'icts where Elk ai'e ahun- 
dant the ti'ee is constantly eaten down, so that it seems likely to 
become much scarcer than at present. 

Scotch-fir twigs are also largely consumed during the winter, 
but do not seem to be eaten so long as mountain-ash can be 
easily procured. Though I have never actually seen Elk editing 
grass, I am assured by the native hunters and by Oapt. Ferrand 
that they do so in sunnner to some extent, and also ])ite oft' the 
fiower-heads of Epilobium and other plants. Water-lily roots, 

* "Saeter" is the Norsk tcini for a shieling. 

2 Capt. Ferrand has known a bull Elk beat oil' the attack of two bears in company. 

138 MR. H. J. ELWES ON [Feb. 3, 

which are said to be the favourite summer food of the Moose in 
America, are also eaten, though the plant is rare in the districts 
where I have hunted. 

The Elk seems to spend the greater part of the day in feeding, 
though it lies down for some hours to chew the cud. Some 
hunters say that it has a regular time for lying down, and 
will not hunt between 11 and 2 o'clock, because of the great 
difficulty of approaching the animal when it is resting. I have, 
however, seen them lying at all hours between 9 a.m. and 3 p.m., 
and have found them up and feeding at midday and later, so that 
there is evidently no rule beyond the appetite of the individual 

From its great size and the nature of its food, the Elk requires 
a much larger extent of feeding-ground than any animal I am 
acquainted with, except the elephant ; and I should suppose that 
at least thi'ee square miles of suitable forest would not support 
more than one Elk continuously, judging from the number I have 
found in places where they were at home and undisturbed. 

It is an animal of extremely solitary habits, and in summer 
more than three will hardly ever be found together, and more 
often only two. A family-party usually consists of a cow, a calf, 
and a yearling ; very often the cow and the calf are alone, and 
two bulls are frequently found together before the rvitting- season 
begins. In winter, however, they are said to be somewhat more 
gregarious, but I have never heard of more than nine actually 
being seen in company in the autumn. 

In some Swedish forests, which are strictly preserved for shooting, 
it is said, however, that the Elk associates in larger herds, and has 
become so numei'ous that much damage is done to the forest by 
their biting off the shoots and tops of the young Scotch pines ; but 
in such localities they do not attain the size and vigour that they 
do whei-e they have a wider range and a greater choice of food, as 
in the two pi-ovinces where I have observed them. 

The Elk is commonly supposed to be essentially an inhabitant 
of forests, and though this is to some extent the case, and in winter 
it no doubt almost invariably keeps to the shelter of the forest, 
yet I have lately become convinced that it is found on the higher 
fjelds to a much greater extent than is generally known. As 
high as the bii-ch is found, Elk may certainly be seen in summer ; 
and the old solitary bulls in disti'icts where open fjelds exist 
rarely descend into the pine-forest until the rutting-season begins. 
From 1500 to 2500 feet is a very common range in summer, and 
on the high mountains of Upper Tydal I have seen tracks much 
higher than this, far out in the open f jeld on the Swedish frontier. 
And in North Ti-ondhjem, where the timber-line is not so high, the 
open hill- tops in some places are covered with Elk tracks and dung, 
as though the animals had remained there dui'ing the whole of 
the hot weather. Capt. Feri'and assures me that he has killed a 
bull on the mountain above Lake Fsemen at an elevation of at least 
3700 feet, on regular reindeer-ground, whei-e arctic willows were 

1903.] THE ELK IN NORWAY. 139 

the only shrubby plants. When found in such situations, they 
may be stalked in exactly the same way a.s led deer, but when 
disturlied usually make for the sheltei- of tlie forest ; and even on 
the fjeld they nearly always lie in or to a small grove of 
dwarf birch, and are in consequence not so e;xsy to find with the 
telescope as red deer. 

The Elk is supposed to be monogamous, and I have never seen 
moi'e than one cow in company with a bull during the rutting- 
senson ; but I am assured l)y a Lapp, whose knowledge of their 
habits is very great, that in districts wheie cows are more 
numerous than bulls, the older bulls will change their mates two 
or thi'ee times diu'ing the season. This seems to be confirmed by 
the fact, which 1 have often noticed, that at the end of Septend)er, 
when the I'utting-season is approaching its height, cows are often 
found without bulls, and the ti'acks of solitaiy Ijulls are found, 
travelling presuma,l)ly in search of fresh cows. 

Col. Walker, of Tykillen, Co. Wexfoixl, who has had great experi- 
ence in Elk-hunting, relates the following event in a letter to me : — 

"Thomas and I saw a magnificent bull with a splendid head in 
company with a cow and calf ; they were quite in the open on the 
edge of a small lake about half a mile from us. 

" I noticed that whenevei' the bull went to the cow she ran at 
him, and butted him very hard in the ribs ; he then each time ran 
from hei', and browsed on the ti'ees. Thomas then told me that 
it would be useless to stalk this bull as he would be gone in a few 
minutes. His stoiy was that the bull only remains -with the cow 
for three days, and then she lieats him oft', and he has to go and find 
another. Thomas had hai'dly done telling me this when the bull 
made his last appeal to the cow. She gave him a rough recejition, 
and he at once stai'ted oft" at a fast trot. She remained grazing, 
and we could seethel)ull going away very fast for nearly two miles 
over a long stretch of open mountain, which took him aci'oss our 
boundary. We never saw this bull again." 

Another pi-oof of their polygamous habit is that on such ground 
as that on which I hunted last year, whei-e duiing eight yeai's only 
one cow has been killed for every five or six bulls, and where in 
consequence the cows were far more numeious, I found no more 
barren cows than in districts hunted by Noi'wegians only, where 
more cows than bulls had been killed. 

A point upon which I have never lieen able to get any certain 
information in Norway is whether the sexes atti'act each other 
by calling at night, as the American Moose do. The art of calling 
is unknown to Scandinavian huntei's so far as I can learn, though 
it is said to be practised in North Russia and Kurland. The 
Aveather in NorAvay is usually so wet and stormy at the end of 
September that thei'e is no inducement to lie out in the forest ; 
and though I have spent many nights in remotely situated huts 
and sjcteis, and have listened at dark and daylight, I have never 
heard such a cjill as is made by the Moose. The cows do, however, 
call to theii' calves, and I have heard bulls uttering a low grunting 

140 MR. H. J. ELWES ON [Feb. 3, 

noise when alone, and apparently looking for cows. On this 
point, howevei', I quote Capt. FeiTand's expei'ience, which is that 
he has on several occasions attracted bulls whose whereabouts 
he knew, especially on a still fi-osty evening, by imitating the 
grunting of the bull, which can be heard when they are looking 
for cows at a distance of from 500 to 1000 yards, or possibly 
moi-e. Moreover, he has heard the cows make a somewhat 
similar noise, and the calves make a bleating noise when they 
have lost their dam. 

Col. Walker also writes to me on the subject of the cow calling 
for the bull, as obsei'ved by him at Storvand, on the property of 
Mr. Oollett near Mo, in North Trondhj em, in 1890 :— 

" One day my hunter and I came on the spor of a bull Elk and 
cow ; after some time we looked over a small hill and saw, in the 
valley below it, the bull and cow about 150 yards from us. Seeing 
that he was paying hei- great attention I waited to see what would 
happen ; after one or two attempts he jumped on her, they having 
their backs to me : I aimed at him, but waited until he had finished 
his performance ; as he came ofi' her I fired, and shot him stone 
dead. He rolled ofi' on to the gi'ound. She did not appear to 
have noticed the shot or that he was dead, but began to graze 
close to him, and once or twice went over and smelt him, then 
grazed again. I then walked up to the bull ; she did not seem to 
mind me, only stai-ed at me and stood close to the bull. When I 
was within twenty yards of her she got alai'nied and went ofi", 
very slowly at first, constantly looking round for him to follow 
her. Shortly afterwards, while we were cutting up the bull, we 
heard a very loud harsh roaring noise, just like the noise made by 
a badly wounded bear when pinned in a corner. Both my hunter 
and I thought it was a bear. I then went to the top of a small 
hill, and on looking over saw the cow in the middle of a large open 
bog. She was roaring, I could see her quite plainly thi'ough my 
glasses ; she then got wind of me and bolted off. 

" On another occasion I had killed a bull which was in company 
with a cow and two calves ; these bolted, the cow one way, calves 
the other. About 7.30 p.m., when I was in my hut, I heard a cow 
calling just like a domestic animal in this country. I asked 
Thomas whether we were near a farm ; he then told me it was 
the Elk cow calling for her calves." 

Though the scent and hearing of the Elk are unusually acvite, 
their sight seems to be by no means so quick as that of deer : if it 
were, the difiiculty of shooting them, which is already veiy great, 
would be much increased. The usual method adopted in Norway is 
to use a trained dog in a leash, which can scent the Elk at a distance 
of a mile or more in the forest ; but owing to the difficulty of 
approaching without noise in the thick forest, and the cunning 
which the Elk almost invariably display in lying down so as to 
get the wind of anyone following their tracks, only those who have 
great patience, caution, perseverance, and an intimate knowledge 
of the habits of the Elk and the ground, are successful in 

1903.] THE ELK IX KORWAY. 141 

getting- more than a glimpse of a frightened Elk disappearing 
among the trees. But if you can succeed in seeing the Elk l){>fore 
he has hea,i-d oi- smelt you, you may take lihei-ties with his eye- 
sight which would not he possihlc in deerstalking. I once 
appi'oached a cow Elk, which was lying with her calf on an open 
hillside, within 10 yai-ds hy keej)ing her hody hetween me and 
her head, and might have got nearer if the calf had not seen me. 
On another occasion I rowed in a boat, just as it was getting 
daylight, within 150 yards of a hull Elk, whii-h had hcon lying on 
the shore of a lake, and had not yet got up from his laii-, and 
shot him from tlie boat before he had taken alarni. 

When bulls are rutting they sometimes run towards a man wliom 
they have heard but not seen, thinking that it is another Elk, and 
in conse(pience stories are told of their attacking human beings. 
But these stories will rai-ely bear investigation, and though there 
ai'e undoubted instances of wounded Elk attacking human beings, 
I never but once saw one turn upon me, and then only because 
he could not get away. 

Besides the system of hunting tbem witli a- dog in a leash, they 
are also hunted to some extent with loose dogs which bring them 
to bay ; and though this system reipiires a man of great endurance 
in order to follow fast enough to keep the hounds within hearing, 
sometimes for many hours, it is perhajis more exciting and deadly 
than still hunting. When Elk are so disturbeil by dogs and 
have not seen, heai-d, or smelt man, they will sometimes come to 
bay veiy soon, but more often I'un many miles, and ah\'ays tiy 
to throw the hoinids and hunter out by crossing large rivers or 
lakes. The Elk is an extremely sti-ong and good swimmei-, and 
(piite at home in the watei-, and though he usually ciosses a large 
river where it is shallow, he can, when pi-essed, pass a. I'oai-ing 
torrent, whei-e no boat could live. In the veiy wet autumn of 
1893 I lost several of the bulls I hunted, in consequence of their 
ci-ossing i-ivers which we could neither foi'd nor swim ; when 
severely wounded an Elk iisuall}' takes to water. 

They may sometimes be driven successfully in places wlune the 
groinid is very steep or confined, but as a, rule Elk do not follow 
particular paths so much as most large animals, a.nd can go up and 
down very steep I'avines anil rocky places which would be thouglit 
impossible for so large and heavy an animal. They usually feed 
u]) wind and I'un down wind, but there is no rule about this, and 
after a long hunt they sometimes come back neai'ly to the same 
ground they started from. 

On the whole, I consider Elk among the most difiicult animals 
to hunt that I have ever had experience of, and, even where they 
are (piite numerous, Ihave more than once spent ten days w Ithout 
getting a fair chance at an animal worth shooting. 

The bulls are said by some to shed their horns as early as 
Christmas and not later than Januaiy, as Mr. Meade- Waldo tells 
me they have done in our Gardens. Capt. Ferrand, however, 
luis been infonncd l)y woodcutters who were worl<iiii,r ;dl winter 



[Feb. 3, 





144 MR. H. J. ELWES ON [Feb. 3, 

in the forest, that March is a more usual time, and this seems 
more probable having regard to the time at which the horns of 
other deer are shed. The new ones begin to grow in April 
or May, and are fully developed about the middle of August. In 
September the velvet is rubbed ofF against a young fir-tree, 
which is usually destroyed in the process, and from the middle to 
the end of September the rutting-season commences. At this 
season the bulls have a very strong, rank, musky smell, and 
scrape shallow round holes in the ground like stags' wallows, which 
retain the scent of their urine for some days. Occasionally the 
bulls fight desperately, and I have seen places where the ground 
was torn up over many yards and sprinkled with hair and blood, 
but few hunters have had the good foi'tune to witness such an 

The development of the horns of the Elk in the Nfiinsos district 
of Norway is apparently greater than in South Noi'way and 
Sweden, or in the districts which it still inhabits in East Prussia 
and Kurland, though I cannot speak with the same certainty as 
to Russia. 

On my I'eturn from Siberia, four years ago, I saw semif ossil horns 
of the Elk from the district of Perm, which were larger than 
any I have seen in Europe ; but on visiting the best collections I 
could hear of in St. Petersburg, I saw none which appeared to me 
better than the best Norwegian heads, or so good as the one 
figured (text-fig. 23, p. 143), which is that of a Lithuanian Elk 
in the Branicki Museum at Warsaw. 

The bull calf has no horns the first year. The second year he 
has a small spike on each side, sometimes, but rarely, two spikes, 
and I have seen one on one side and two on the other. As he 
grows older it is supposed that the number of points increase 
annually, one on each side ; but I think this is not at all invariably 
the case, and that the size and strength of the individual has much 
more influence than the a,ge, and probably, as in the case of other 
deer which shed their horns annually, food and climate have a 
good deal to do with it. As a rule the horns increase in size and 
number of points up to 10 or 12 years old and possibly more, and 
in old animals have a tendency to diminish in size and in the 
length and strength of the points. From 18 to 22 points are 
pei'haps the aveivage of adult males in Noi'thern Norway, and 10 
to 14 points in Southern Norway. 

The widest span which I have ever seen is 54 inches (text- 
fig. 18, p. 134), and the greatest number of points 16 on one side, 
as on the shed horn of which I show a tracing (text-fig. 22, p. 142). 
The breadth of the palm in this instance and the numbei' of points 
are quite exceptional, and I very much doubt whether there are 
any such Elk now alive in Norway, as Herr Bruun of Trondhjem, 
who has the opportunity of seeing all the finest specimens which 
are procured, says that he has seen nothing equal to them. 

Moose heads of as great or greater size are, however, often 
killed, and though the largest which I have ever seen is 65 inches 

1003.] THE ELK IN NORWAY. 145 

iicrnss, J liave lieard on t'.iii-ly good ivutlioi-ity of ono ovov (\ foot in 
s|):iii. (This was written before anytliinn' known <>[ iJio 
^i^'aiitie iioiuls found in Alaska.) 

VN^itli regard to tlie comparative size .-uid woiglitof the KIk .ind 
Moose, I cainiot speak positively, as it lias never been |)ossil)le to 
weigh an entire animal in tlie places where I have killed them. 

Ml-. Abel Chapman, however, givo^s me the following weights of 
a bull, of fair but not unusual size, killed by him on Se[)t. liltl) : — 

KilonT.aniincs. ICilosiraiiinics. 

TToad with skin of neck 40 = 40 

Shoulders 42 each = 84 

Ha rinches with foot 54 „ =108 

Si.los 40 „ = 80 

Nock and back 60 = GO 

Skin, say 30 = 30 

Total 402 

This would make the weight of the living animal at least 
1000 pounds or over, as the intestines are enoi'mous. 

Capt. Feriand's best bull, a lO-3'^ear old, weighed 1240 English 
pounds witliout the intestines. 

The pa,ce of Elk, when undisturbed, is a slow walk, and their 
movements are very deliberate, but they can ti-ot for many 
miles over boggy and I'ough ground at a pace of 6 or 8 miles an 
hour, and occasionally when much frightened break into a 
Inmbering canter. 

The female Elk has her first calf at three years old in tlie month 
of May, which malces the pei-iod of gesta,tion about 7 months. 
Usually she has only one, but not unfi-equently two cahos, \\hich 
grow very rapidly, and ))y the end of September are as big as a 
red-deer hind. They suck their dams until late in the winter, 
and keep company with them until another calf is dropped, and 
sometimes longer. They have occasionally been tamed in Sweden 
and taught to go in harness, but owing to the difficulty of feeding 
them they are not easy to keej) in confinement. I ha^-e seen, 
however, in the Zoological Gardens at Rotteitlam a, cow Moose 
which is said to have been about 20 years in confinement, and 
I hope that we shall be equally successful with the one which was 
captured by Mr. Nickalls and presented by him to oui- Gardens. 

Statistics of the numl)ei' of Elk kilk^d in Norway in the season 
of 1894. (Translated from the ' Tidsskrift Norsk. Ja-ger og 
Fisker Forenings,' Heft 3, 1895.) 

Name of All! t. No. of P.iills. No. of Cows. Totnl. 

Smaalenene 8 3 11 

Akershus (w 60 127 

Hedemarkou 74 72 146 

Kristians 90 87 177 

Prog. Zool. Soc— 1903, Vol. I. No. X. 10 



[Feb. 3, 

Name of Amt. 


Jarlsberg og Larvik 


Nedenes , 

Lister og Mandal . , 
Sondre Trondhjems 
Nordre Trondhjems 



Sondre Bergenhus . 
Nordre Bergenhus ., 



No. of Bulls. No. of Cows. 





















646 606 1252 

Average for the 5 years, 1889-93. 610 512-6 1122-6 

To give a better idea of the numbers of Elk found in what was 
perhaps, at that time, the best forest in Norway, I give the 
following statement copied from my diary in 1895. The number 
actually seen may possibly represent from ^ to g of the number 
actually on the ground at the time : — 

Cows & 

Bulls, yearlings. Calves. 

Sept. 2. Haende 1 ... Barren cow shot for meat. 

3. Storen ... 1 1 

4. Strommen... ... 1 1 

5. Strommen... 1 11 ... 1 bull killed : 18 points. 

6. Blank day. 

8. Mo Ill 

9. Mo 1 

10. Oplo 1 1 

11. Syiinses ... 1 

12. Lia Ill 1 

13. Lia 11 ... ... 1 bull killed: 16 points. 

15. Blank day. 

16. Mo Ill 

17. Storengen 1 1 

18. Storengen ... 1 1 1 

19. Fos 1 1 

20. Mo 11 1 

21. To Storvand, blank day. 

22. Vennevik ... 1 11 1 1 bull killed: 6 points. 

23. Synnffis 11 111 1 1 

24. Brotten 11 ... ... 1 bull killed: 14 points. 

25. Synnses 1 1 1 1 bixll killed : 13 points. 

26. Quisten 1 ... ■■- 1 bull killed: 11 points. 

27. Oplo 1 1 

29. Mo Ill 1 bull killed : 18 points. 

Total 17 25 14 

This number of Elk was found on about 12 farms out of 24 
over which I had the right of hunting. These farms might 
average 10,000 acres more or less in extent. 

On the same ground as this Col. Walker kept the following 
record of the Elk which he saw himself during the four seasons 
1887-1890, between Sept. 1st and Oct. 10th. It tends to show 




how rapidly Elk increase where tliey are protected and the cows 
not killed : — 









j 10 
8 7 

17 14 

18 1 20 


Two of tliosc cows hiul two ("lives each. 
One of these cows had two calves. 
Six of these cows had two calves each. 

Nine of tliPS(! cows jiad two calves each., 


Most of these observations were wiitten at the reqnest of 
onr late President, Sir W. Flower, six years ago, but were 
withheld from pnlilication until this year, when the question 
of a new species of Elk existing in Siberia was raised l)y Mr. 
Lydekker and thejion. W. Rothschild. I then added the following 

Text-fio-. 24. 

Morns of a very old Elk showing- deijeneration. (About ^V nat. size.) 

Appendix. — On the Variation op the Elk. 

J)v. Li)nnl)ei'g's papei' on the vai'iation of the Elk appears to 
me to confirm the opinion I expi'essed at a recent meeting of the 
Society (P. Z. S. 1902, vol. ii. p. 144), that the Siberian Elk 
described by Mr. Lydekker (P. Z. S. 1902, vol. i. p. 207) as Alces 
hedfordia' was nothing more than an inconst;int Viiriety of the 
Eui'opean Elk. The Hon. Walter Rothschild has since expre.ssed 
M contrary opinion, foi' which, however, I am not aware that he 
has any fuither evidence than the fact that other horns similar 




[Feb. 3, 

to those of the so-called A. bedfordice have been received by him 
from Sibeiia. I stated that the only horns of the Elk which I 
succeeded in obtaining in the Altai Mountains in 1898, which 
are now in the St. Petersburg Museum, were palmated in pre- 
cisely the same manner as those of the Norwegian Elk, and others 
which 1 have seen in Russian collections had all well-developed 
palmation. Dr. Lbnnberg's figures go to show that the develop- 
ment of the antlers in those parts of Sweden from which they 
come is nothing like so fine as in the districts of North and South 

Text-fig. 25. 

Tracing of a cast horn found at Solem in Bangdal bj' Capt. Ferrand. 
(About ~ nat. size.) 

Trondhjem, where most of my hunting has been done. When I 
first began Elk-hunting in Tydal, a mountain valley in South 
Trondhjems-amt running up to the Swedish frontier, I should have 
considered the horns figured by Lonnberg in fig. 3 as a fair repre- 
sentation of an adult Elk in that district ; but I have seen them with 
as many as eleven points on each horn, and believe that horns with 
even more points have been obtained. Further north, in North 




Ti-ondhjems-amt, wliei-e the Elk a few yeai-s ago was exti-emely 
numerous, mucli lai'gei- heads occurred, owing, I believe, to the much 
greater quantity of mountain-;ish, which seems to be the favourite 
winter food of the Elk, anel wliicli has been to a great extent 
desti'oyed by the vast numbei' of animals constantly devoui'ing it. 
Out of the whole numlier of bulls I have killed, about twenty, 
only four oi' five had horns of the type shown in Lonnberg's 

Text-fiL^ 26. 

One-homed Elk in Ipswich Museum. 

figui'es 1, 4, 6, and 9 ; and all of these, with one exception, 
appeared to be quite young animals. 

Among the huntei's who accompanied me in different years 
were a Swede, a Norwegian, and a Lapp, all of whom knew the 
Elk of those districts most intimately, and were considered the 
most expei-ienced huntei's of theii- districts. None of them ever 
even sviggested the possibility of two races of Elk existing ; and I 

150 ON THE ELK IN NORWAY. [Feb. 3, 

attribute non-pahnation of the horns entirely to impeifect 
development of the animal, caused by insufficient food in their 
earlier years, or degeneration caused by old age, wounds, or other- 
wise. We did not reckon an Elk to be adult until he had at 
least seven points on each horn, and the oldest and largest bull 
that I ever killed, which had been well known in the district, and 
continually hunted for at least ten years, had, when I killed him, 
well-palmated hoi-ns of about one-half the size of what they ought 
to have been in an adult animal of his size, showing that in the 
Elk, as in the red deer, the horns degeneiute in size and number 
of points in old age, which may be fifteen or twenty years or less. 
Two such instances of degeneration are figured. Text-fig. 24, 
p. 147, is the head of an old Elk killed by Thomas Bate, Esq., in 
Lurudal Namdalen on Sept. 25,1 890. The horns measure 43 inches 
in expanse, with ten points on one side and eleven on the othei- ; 
the points are, however, not arranged in a uniform series on the 
edge of the palm, and the development of the brow- antlers is very 

The shed hoi'us of what was probably the same animal were 
picked up on the same ground in the year previous by Col. Sullivan, 
and ai'e of the same type with the same number of points. 

An Elk displaying still more remarkable abnormal degeneration 
was killed by Capt. Ferrand, and is shown in text-fig. 26, p. 149. 
This animal was supposed to be 25 years old or more, aiid 
the incidents of his death have been most graphically described in 
the ' Badminton Magazine ' for March 1901 by Caj)t. Ferrand. The 
horns are now in tlie Ipswich Museum. 

The largest horns I have seen from Norway, belonging to an 
animal which I unsuccessfully hunted for many days, but which 
was afterwards killed by a farmer, and sold to me by Mr. Bruun 
of Trondhjem, were 54 inches in width, with nine points on each 
side (see text-fig. 18, p. 134); but there is a pair of shed horns in 
Sir Henry Pottinger's house at Mo, of one of which I send a tiucing 
(see text-fig. 22, p. 142), showing sixteen jooints on each side. It 
is well known in Norway that the Elk of the southei'n districts, 
which are much more fully timbered, and whei'e there is nothing 
like the same extent of open fell and good feed as in North 
and South Trondhjem, and where both sexes are much more 
constantly hunted and the calves frequently deprived of their 
mother's milk in Sej)tember, do not, in modern times at least, 
produce anything like such fine heads as those of the wilder 
districts of the north, the conditions being probably very similar to 
those described by Di-. Lonnberg as in the southern provinces 
of Sweden \ Taking his nine figures, I should be inclined to say 
that all except 2 and 3 might, if they had come from Siberia, 
have been considered as belonging to Alces bedfoj-dice ; and form, 
to my mind, ample proof of that being (if the horns belong to 

1 Mr. Percy Godman informs me that a well-known Elk-hunter in South Norway 
considers that on his propertj' Elk have diminished by two-thirds in the last ten 
years, and attributes this decrease to their having eaten and destroyed all the " leaf- 
trees," i. e. willow, mountain-ash, and aspen. 

P.Z.S. 1903, vol. 1. pi.xiy. 

J.Picltarfl Gambridle (tfil etlith. West,NewiTian imp, 


P.Z .S. 1903. vol.1. PI. XV. 

^""""'^ 17. 
F. Pic:Kard.Ca.mbri(ige.(lel.tfditii. WG5t,Newman imp. 



adult animals) iiotliiiig more than a local variety produced by com- 
pai'atively unfavourable conditions. In consideiing the prol^ability 
of the existence of local races of Elk, it must be remembered 
that no northern mammal, except peihaps the reindeer, has such 
wandering habits ; and though a race might })ecome temporarily 
isolated in an area, which was entirely sui-rounded by a lai'ge 
extent of country providing no suitable food, yet there is no 
doubt, in my mind, that when their favourite food had become 
partially exhausted, they would migrate many hundreds of miles, 
and thus pi'event the establishment of local races. 

A very interesting piper on the former existence of the Elk in 
the Thames Valley in England, with a plate showing the great 
similarity of its horns to those now existing, is j^iiblished by 
Mr. E. T. Newton, F.Ii.S., in the Quarterly Journal of the 
Geological .Society, vol. lix. (1903), and I am much indebted to 
that gentleman for sending it to me. 

February 17, 1903. 

Dr. Henry Woodward, F.R.S., Vice-President, 
in the Chaii-. 

]Mr. R. E. Holding exhibited and made remarks upon an adult 
skull of a Collie Dog, indicating a displacement of the incisors 
caused by the closing of the left lower canine upon the third 
incisor of upper jaw, and which apparently retarded the develop- 
ment of the second incisor of that side. The skull also showed a 
supernumerary canine which could in no way be mistaken for a 
retained milk-canine. 

Mr. Holding also exhibited portions of three Rabbits' skulls 
having "over-shot" incisor teeth. In one specimen the abnor- 
mality had caused a deviation fi'om the median line of the anterior 
portion of the skull ; and in another the incisors, after leaving the 
pi-emaxilla, had formed a complete circle and grown into the palate, 
causing stai'vation and death of the animal. 

The following papers were read : — 

1. On some new Species o£ Spiders belonging to the 
Families Pimuridcc and Senoculichc ; with Characters of 
a new Genus. By Frederick Pickard-Cambridge, 
B.A., F.Z.S. 

[llccciveil Jamiary 10, 1903.] 

(Plates XIV. & XV.') 

While working out the Spiders of the two families Pisauridm 
and HenocuUdai of Central America for the ' Biologia,' oppor- 
tunity was kindly offered l)y Mr. R. I. Pocock for an examination 
of the material in the British Museum Collection, with the result 

1 For u.\plaiiatiou of the Plates, see p. 168. 


that a new genus and eight new species were found, several of 
them occuiring amongst those collected by myself on the Lower 
Amazons in 1895-6. 

The Spiders belonging to the Fisaimdce are interesting from 
the fact that they run freely and with great rapidity over the 
surface of the watei", even in a sti'ong curi-ent. One of the genera, 
jDolomedes, and another, Thalassius^ have both been dredited with 
captiu'ing and devouring small fish of various kinds ; and of the 
latter genus Mr. A. N. Stenning, himself formerly a gamekeeper, 
whose observations are likel}^ to be trustworthy, declares that he 
has found members actually devouring the small fry of a variety 
of trout which occurs in South Afi-ica, some of the culprits being 
now in the Museum Collection. Other notes on the habits of 
these intei'esting Spiders will be fovmd under the genus Trechalea. 

Species descrihed or figured heloiv. 

Thaumasia velox B. Simon, 5 > P- 154, PL XIV. fig.' 1. 

,, anmdipes, sp. nov., 2 ) P- 154, PI. XIV. fig. 2. 

Dossenus marginatus E. Simon, 5 , p. 155, PI. XIV. figs. 3-5. 
Paradossenus nig?'icans, sp. nov., $ , p. 155, PI. XIV. figs. 6-9. 
Thanatidius spinipes, sp. nov., 5 , p. 156, PI. XIV. figs. 10-12. 
/Senoculus pcmrdlelus E. Simon, $ , p. 166, PI. XIV. fig. 13. 

cdbidus, sp. nov., 5 , p. 168, PI. XIV. fig. 14. 
Trechcdea longitarsis 0. L. Koch, c5" 5 , p. 160, PI. XV. figs. 13-17. 
„ keyserlingi^ sp. nov., 5 ? P- 163, PI. XV. figs. 1, 2. 
,, urinator E. Simon, J $ , p. 161, PI. XV. figs. 3--5«. 
,, ellacombei, sp. nov., 5 , p. 161, PI. XV. fig. 6. 
,, macconnelli Poc, (^ , p. 162, PI. XV. figs. 7, 8. 

coniiexa O. P.-Oambr., d , p. 162, PL XV. figs. 9, 10. 

extensa 0. P.-Oambr., c^ , p. 162, PL XV. fig's. 11, 12. 

,, amazonica, sp. nov., J $ , p. 163, PL XV. figs. 18-20. 

Hesydrus pahistris E. Simon, (S $ , P- 165, PL XV. figs. 22-25. 

hahilis O. P.-Oambr., S , p. 165, PL XV. fig. 21. 

Fam. PisAURiD^. 

/Synopsis of Genera decdt loith below. 

A. Tibia i. with four (2-2-2-2) or five (2-2-2-2-2) 
paired spines beneath, the last pair very small and 
apical. Protarsi i. with three or four pairs beneath. 
A'. Tibia i. with four pairs of spines beneath. Protarsi 
i.-iv. with a single small central apical spine beneath. 
Sternum produced into a long narrow conical point 
between coxaj iv. 
i. Tarsi shorter, not flexible. Protarsi i. with three 
(2-2-2) paired spines beneath. 
aa. Lower margin of fang-groove with 3 teeth. 
a. Tibial and protarsal spines comjiaratively short, 
only two or three times as long as the diameter 
of the segment. 
* Anterior row of eyes wider than that formed 
by the posterior centrals. Central posterior 
eyes nearer to each other than to the 
a^. Teeth on lower margin of fang-groove not 

equidistant, the 3rd more remote Thaumasia Perty. 




6'. Teuth on lower margin of fang-groovu 

(■i|ui(list:int Tiniis V. I'.-Caml.r. 

** Aiitfvior row of eyes not wider than that 
fornii'd ])y the posterior centrals. Cen- 
tral posterior eyes much further ai)art 

than each is from tlie lateral Enna O. I'.-L'amljr. 

b. Tiliial ami jirotarsal sjiiiies com])arative]y 
very lon.y;, the lon;,'-est heini; ([uite two 

thirds tlie length of the whole segment .. JJossemis E. Simon. 
////. Lower margin of iang-groove with 4 teeth, 

the 3rd smaller I'aradossenus, gc]\. nov. 

ii. Tarsi long and ilexihle. Protarsi i. with four 
(2-2-2-2) i)aired si)ines heneath. 
a*. Leg.s sul)ei|ual in pairs, l — 2 and 1 — 3. .Man- 
dibles conspicuously gibhous ahove, glahrous, 
more distinctly earinate on the outer side 
(males). Lower margin of fang-groove with 

3 teeth Uesjjdrus Simon. 

b*. Legs unequal, 4, 2, 1, 3. Mandibles not 
conspicuously gibbous above, hairy, less 
earinate (males). Lower margin of fang- 
groove with 3, d, or 5 teeth Trccliaha Thorell. 

H'. 'I'iliia i. with five jiairs of spines beneath. Pro- 
tarsi i.-iv. with two apical spines beneath. 
Sternum very broad between co.xa; iv., not 

forming a point 'i TliauaLidius K. Simon. 

1>. Tibia i. with 9-10 pairs of spines beneath, the last 
pair not ai)ical. Protarsi i. with 8 or 9 pairs of 
spines beneath Sijidrechalea F. P.-Camlir. 

If the above genera he separated l)y the perhiips moi'e I'asily 
observed character furnished hy the eyc-fornuda., then the tal)le 
would run as follows : — 

A. Anterior row of eyes not or only a little wider than the posterior 

central row. r Enna 

a. Anterior row of eyes not wider than the posterior central row... \ jjossenus 
h. Anterior row of eyes a little, but distinctly, wider than the 

posterior central row Faradossemis. Timis. Thaumnsia. lles,i/dri(s. 

Trcchalea. Sijntrcchalca. 

B. Anterior row of eyes much wider than the posterior central row, 

as wide as the posterior lateral row Thanalldius. 

Note. — (1) The number of protarsal sjiines in some species of Trechalea arc 
difficult to locate, and the character is thus rendered less valunlile. 

(2) One cannot be ([uitc certain that Thaii<((idiiis has lieen correctly identified. 

(3) Tiiins is closely allied to Dranccs Simon, but the latter has, according to this 
author, no apical pair of spines beneath tibia i. and ii., while they are present in the 
former. The name Urauces (not Dranccs) was used by Champion for Coleoptcra 
in 1889. 

(1) In the l?iologia Centr.-Amer., Arach. Aran. ii. i). 312, I have referred the 
species Hcsi/drus juUicni Simon to Enna 0. P.-Cambr. on the ground that they 
have the characters, the tarsi flexible and central jjosterior eyes 2 diameters apart, in 
common. This geims therefore comprises Simon's JJcsj/dnis Sect. 2 a, while 
Hest/drus, as here diagnosed, comprises bis Jlcsi/drus Sect. la. 

Genus Tiiaumasia Perty. 
Del. Anini. Art. Bras. p. 192 (1833). Type, T. senilis Perty. 

SjJecies recorded. 

1 . T. senilis Perty, loc. cit Brazil. 

2. T. nuinjinella (C. L. Koch) (Sub JJolomedes.) 


3. T. sGO'paria (E. Simon) (Sub Scdtuinus.) 

4. ? T. scajndaris (0. L. Koch) (Sub JDolomedes.) 

5. ? T- binotata (0. L. Koch) (Sub Dolomedes.) 

6. Thaumasia velox Simon, Ann. Soc. Ent. Belg. xlii. p. 18 

(1898) Brazil. 

Thaumasia velox Simon. (Plate XIY. fig. 1 .) 

Loc. cit. Type S , in coll. E. Simon. Total length 8 mm. 

Thaumasia velox F. P.-Cambr. Biol. Centr.-Amer., Arach. Ai-an. 
ii. p. 309, t. 30. figs. 5, 5 a-c ( 6 )■ 

The type of this species {<S) was taken on the Amazons, and, 
short of first-hand evidence, I do not doubt that the male and 
female, taken by myself amongst the foliage on the margin of a 
small " furo" in the Parana Buyassu, near Bi'eves, Lower Amazons, 
must be referred to this species. 

The male identified as belonging to this species from Guate- 
mala may possibly be not identical, but it is impossible to be 
certain with but one example from each locality. 

The female has not been before described, but a figure of the 
vulva is given on the Plate. For figures of the male see Biol. 
Centr.-Amer. loc. cit. above. 

Hah. Lower Amazons : Santarem ; Breves, Parana Buyassu 
{F. P.-C, 1895-96); Rio Tocantins, S. Paulo de 01iven9a {de 
Mathan) ; Guatemala (Sar-g). 

7. Thaumasia annulipes, sp. nov., $ . (Plate XIV. fig. 2.) 

Type 2 , in coll. Brit. Mus. Total length 5-5 mm. 

5 . — Colour. Carapace deep bi'own, with pale yellow nai'row 
central band, slightly dilate behind eyes. Abdomen brown, 
shoulders and anterioi' lateral portion deep black. Legs pale 
yellow, femora annulated with dusky brown. 

Structure. Carapace horizontal above, convex and abruptly 
deflected behind. Eyes : posterior row recurved, subequal, equi- 
distant ; latei-als on a stout black tubercle. Anterior I'ow straight 
or slightly procurved, wider than width occupied by jDosteiior cen- 
trals. Anterior eyes equidistant, centrals larger. Clypeus more 
than three tiraes the height of the diameter of anterior centrals. 
Mandibles long and stout. Fang-groove with three denticles on both 
margins. Sternum cordiform, posteiior angle attenuate, produced, 
its apex extending as far as the pedicle. Coxce of pedipalp long, 
straight, slightly dilate and i-ounded at apex. Lahmm half the 
length of maxillfe, broad, squarely truncate at apex. Legs 
4, 1, 2, 3 ; iii. the smallest, extending as far as apex of tibia iv. 
Tibia i. and ii. with 2-2-2 spines beneath ; 1 — 1 lateral on apical 
half ; 1 — 1 at base and apex above ; 2 at apex beneath. Pro- 
tarsus i. and ii. with 2-2-2 spines beneath and 1 at apex ; 1 — 1 
on each side ; none above. For vulva, see PI. XIY. fig. 2. 

Several females of this small Spider were taken by myself in the 
neighbourhood of Manaos, Lower Amazons, 1895-6. 


(lenus DossENUS Simon. 

Hist. Nat. Ar. 2, ii. p. 314. Type, D. marylaatas E. Siuiuu. 

Lowei" Amazons. 

DossENUs MARcaxATtis E. Siiiion. (Plate XIV. figs. 3-5.) 

Ann. Soc. Ent. Belg. xlii. 1898, p. 19. Type $ , and androtype 
<S in coll. E. 8imon. Total length, 5 , 7"7 mm. 

Two adult females of a Spider, wliich I feel pretty certain nnist 
l»e i-efei'i-i'd to this species, wei'C taken by myself at lireves and 
Parti, and are now in the British Museum Collection. The sj lines 
on the legs are very long in compai'ison with those of allied 

Hah. IsL. Tui.viDAL) ; BitAzir. : Ama/ons; Matto (Jrosso {Ger- 
main); Lower Amazons, Pani, Breves (Parana Buyassu, /'. l\-<'., 

Paradossenus, gen. nov. 

Type, P. nigricans, sp. n. Lower Amazons. 

Lower margin of fang-groove with 4 teeth, the 3rd being 
smaller. Tarsi shoit, straight, not flexible. Pi-otarsi i. with 
3 paired spines beneath ; protarsi of all fom- pairs with a single 
central apical spine beneath. Til)ia i. with four paired spines 
beneath, the last pair apical. Sternum produced into a conical 
point between coxaj iv. Spines on legs short, two to three times 
the diameter of the segment. Posterior row of eyes recurved ; 
eyes subequal, centi-als one diameter apart, two and a half from 
the laterals, which are set on a low tuljercle. Anterior row 
straight or very slightly procurved, slightly wider than the central 
posterior row. Clyjieus one and a half times the height of the 
diameter of an anterior central eye. 

Legs 1, 2, 4, 3 — iii. very shoi't, extending as far as two-thirds 
of tibia iv. 

Paradossenus nigricans, sp. nov. (Plate XIV". figs. 6-9.) 

Type S J 8 mm. ; gynetype $ , 8"5 mm. — carap. 3*75 mm. 
In coll. Brit. Mus. 

d 2 . — Colour. Carapace dull yellow-lirown, with two longi- 
tudinal dark brown bars on each side of the pale central line, the 
whole lieing almost entirely clothed with yellow-grey scale-like 
hairs. Margins of cai'apace i-eticulated with fine dusky lines. 
The anterior })ortion of the brown bars presents within their 
margin a narrow elongate accent-like pale mark. Abdomen pale 
yellow ; dorsal area with a broad longitudinal dark brown (some- 
times paler) foliated band, becoming narrowed towards the 
spinners, from the centre of whose lateral margin, on l)oth sides, 
issues an ol)licpie interrupted lateral bar. 'I'he centre of the 
foliated band is occupied by ,a pale lanceolate liand, much attenu- 
ated towards the spinners, its basal portion again occupied by a 
broad lanceolate dark band, scarcely I'eaehing the centre of the 


abdomen. Lateral ai-ea mottled with dusky brown ; ventral 
sui'face pale yellow, unicolorous. Legs entirely yellow : i. and ii. 
very faintly annulated and speckled with dusky brown, iii. and iv. 
annulated, in some cases very richly, with black or brown. 
Mandibles i-ed-yellow, with a large brown patch on each in front. 

N.B. — These coloui's are very variable and in some cases the 
central dark band within the inner pale band on the abdomen 
is itself pale. 

The vidva consists of two parallel plates, which have a promi- 
nent angle on their inner margin. Between these, in front, lies 
a deep transverse semicircular depression, and a convex central 
piece, with a few shallow longitudinal grooves, is situated between 
the plates. 

For figui-es of tibial spur of male palpijs and the vulva, see 
Plate Xiy. figs. 7, 9. 

These spiders are very abundant on the Lower Amazons, racing 
over the surface of the water with legs extended and body flat, 
and are exceedingly difficult to catch. None of the females had 
egg-cocoons with them, but they pi'obably cany them in the same 
position as Trechalea, for in habits and general appearance these 
spiders ai'e very similar to those belonging to the latter genus. 

Hah. Lower Amazons: Breves, Parana Buyassu {F. P.-C, 

Genus Thanatidius E. Simon. 

Hist. Nat. Ar. 2, ii. p. 293. Type, T. dubius (Hentz). 

N. America. 

The following thi'ee species ai'e referi'ed to this genus by 
Simon : — 

1. T. dubius (Hentz), Bost. J. N. H. v. p. 449 (1847). 

(Sub Thomisus.) 

2. T. tcenius (Hentz), loc. cit (Sub Thomisus.) 

3. T. imdulatios (Keys.), Yerh. z.-b. Ges. Wien, p. 486 (1887). 

(Sub Tetragonophthalma.) 

4. Thanatidius spinipes, sp. nov. (Plate XIV. figs. 10-12.) 

Type 5 in coll. Brit. Mus. Total length 6 mm. 

Hab. Parana Buyassu, Breves, Lower Amazons. 

5 . — Colour. Carapace, mandibles, sternum, and legs bright, 
unicolorous orange. Abdomen olive-green, with a central band 
of cretaceovis- white pigment-cells. 

Structure. Carapace nearly circular, ocular region produced, 
parallel- sided, slightly raised. Thoracic region gibbous, abruptly 
deflected behind. Eyes : posterior row strongly recurved, eyes 
subequal, less than one diameter apart, equidistant ; laterals on a 
stovit conical tubercle. Anterior row strongly procurved ; eye- 
tubercles subequal, less than one diameter apart, equidistant, much 
wider than the width occupied by lateral posteriors. Glypeus., 
in centre as high as four diameters of anterior central eyes. 


Mandibles long, slender, more than twice as long as lieiglit of 
clypens. Famj-groove with tln-ee dentieiilos on lower uiaigiii. 
MaxilliH long, parallel, dilate at apex; lal)imii Iiroad, liall' 1l:c 
lengtli of maxilhe, ti'uneate at apex. 

Sternum longer than broad, cordifonn, produced ])Ostoriorly 
between coxai iv. into a hroad rounded j^lfote, the coxa^ being their 
own diameter apart. Leys 1,4,2,3. Femora with two long 
doi'sal spines and seveiul othei's on the sides and apex. Femnr i. 
with two long spines in front in apical half. Ti])ia i. and ii. with 
2-2-2-2-2 long spines beneath, the longest (the basal) being half 
as long as the segment itself. 1 — 1 lateral spines ; 1 — 1 dorsal. 
Protarsus i. and ii. with 2-2-2-2 long spines beneath (basal [)air 
longest) : 1 — 1 lateral, 1 — 1 dorsal. Spines on tibia and pro- 
tarsus iii. and iv. numerous, but less regulai-. Patella) 1-4 with 
a single long apical spine. Scopula absent. Tarsal claws 3, 
superiors with thi'ee or four denticules beneath. Spinners 6, 
postei'iox'S longest. Pedipcdp with large tarsal claw, having 3 
very long denticules beneath towai-ds l)ase. Vulva, see PI. XIY. 
fig. 12. ■ 

Hab. A single 2 fi'oni the Paiuna Buyassu, near Breves, Lower 

Genus Treciialea Thorell, 18C9 (nom. nov. for Triclaria). 

Type, T. longitarsis (0. L. Koch), Colombia. Sub Triclaria 
(1848), nom. pra>occ. by Wagler for Aves (1832). 

The following characters are common to all the species men- 
tioned below : — Teeth on uppei- margin of faiig-groove 3, on the 
lower, 3, 4, or 5. Tarsi long and flexible. Tibia i. with four 
pairs of spines beneath, the last pair small a,nd apical, besides two 
latei'al spines. Protai-si i. with foui' paii-s of spines beneath (often 
ii'regularly situated) ; protarsi of all foiu' pairs with a small 
central apical spine beneath. Anterior I'ow of eyes slightly 
I'ecurved, extending laterally slightly beyond the posterior 
centi'als. Lateral antei-iors smaller than the central antei'iors. 
Legs variable in length, 4, 2, 1, 3. Clypeus varialile in height, 
porrect, and the eyes also vai'iable in their i-elative positions. 

As regards the type-species of the genus, two distinct forms 
have been identified as longitarsis of Koch — one by Keyserling, 
now in coll. Biit. Mus., a female, having 5 teeth ; the other by 
Simon (Ann. Soc. Ent. Belg. xlii. p. 20, 1898), male and female, 
having 3 teeth. Since they cannot both be the true longitarsis, 
I describe Keyserling's identification as a new species, for it is 
totally distinct from examples from Colombia, now before me, 
whence the t)/pe of the genus originally came. It forms, indeed, 
the type of a, distinct grou]), so entirely difterent is tlie form nf 
the vulva from that of the typical TrecJialea from Colombi.i. 

At pi'esent, how'ever, although the identity of T. longiiarais 
cannot be settled with absolute certainty until an examination 
of the ty})e-specitii('n (if it slill exists in 1lii' Imperial Mnsenm at 


Berlin) has been made, the examples from Colombia are regarded 
and here described as T. longitarsis C. L. Koch. All of the 
species described and recorded below seem to be good and distinct, 
though of course much more material from all parts is required 
for comparison and the confirmation of characters. 

The habits of these spiders are very interesting, for although 
they are not amphibious in tlie true sense of the term, as are 
Argyroneta and Desis, they are quite at home on the water, a.nd 
speed along over the surface with great ease and rapidity, the 
hairy clothing of the legs being waterproof. When racing away 
over the water, they are not easy to distingviish from the large 
Hemipterons (probably a species of Ranatra) of similar habit, 
when the two creatures happen to be together in the same locality. 
On being pursued over the water, the spiders will rush up the sides 
of the boat and take refuge under the boards, but I have never 
observed them dive below the surface as do Pirata, Dolomedes, 
and some other Pisauridte. Whether, as has been reported of 
certain species of Dolomedes and Thalassius, the spiders actually 
prey upon small fish of various kinds, I cannot say from actual 
experience, though the long flexible legs and exceedingly sharp 
claws are well adapted for the piu'suit and capture of such pi'ey 
in their native element. I may add that McOook's records of the 
fish-catching capacity of Dolomedes (Amer. Spid. vol. i. p. 236 and 
iii. p. 66) has recently received confirmation by Mr. A. N. Stenning 
in South Africa. He tells us that Thalassius, a genus represent- 
ing Dolomedes in the Ethiopian Region, has been often observed 
by himself in the act of devouring the small fiy of a species of 
trout, and calls the attention of pisciculturists in those regions to 
the fact, and begs them to keep an eye on these spiders. 

On a swampy island in the lai-ge largo opposite Santarem on 
the Amazons, I have taken examples of one beautiful species 
squatting flat, like Sparassids, on the truidvs of the trees, where 
their hoary-white hairs and mottled legs and body afibrded them 
excellent protective colouring. Nor do they resemble Heteropods 
meiely when at rest, foi- when disturbed they dash round to the 
opposite side of the tree-trunk with all the rapidity of a Selenops. 
The egg-cocoon, as in many Pisaurids, is carried by the female 
attached to the central pair of spinners at the tail-end of the 
abdomen ; and it is cui'ious to note how, when these females stop 
suddenly in headlong flight, the weight of the cocoon swings them 
round on the watei-, and carries them along backwards for some 

The species recorded and described below may be recognized by 
the following charactei's : ^ 


A. Lower margin of fang-groove with 4 teeth, the third 

smaller. Legs unicolorous urinator E. Simon. 



B. Lower margin of f;inp:-jiroove with 3 subequal teeth. 
Legs more or less si)otted or annulated. 

I. Legs longer in ])roportion, leg iv. being about 8 

times longer than the carapace. 

a. Third tooth on U)wer margin of fang-groove more 

remote from the others. 

rt'. Tibia of pedipalp fromoue-half to twice longer 

than broad ; tibial spur distinctlj' unculate at 

apex, not concave on inner side. Embolus of 

bulb with a long conspicuous spur at its liase. longitarsis C. L. Koeli 
/>'. Tiliia of pedipalp not longer than broad ; tibial (sec. F. I'.-C). 
spur not unculate at a])ex, concave on inner 
side, the inner margin having a prominent 
angle. Embolus of bulb without any long spur, 

but having two chitinous ridges macconnelli Pocock. 

h. Third tooth on lower margin of fang-groove not 
remote, the 3 teeth almost e(|uidistant. 
n-. Tibial spur not unculate at the apex. 

Endjolus of bulb without any long spur connexa 0. P.-Cambr. 

h'-. Tibial spur unculate at its apex. Embolus 

of Inilb with a long spur at its base extensa O. P.-Cambr. 

II. Legs shorter in proportion, leg iv. being 5-6 times 

longer than the carapace amazoniea, sp. n. 

An alternative table for Males. 

A. Embolus of bull) with a long conspicuous erect black 

spin- in its basal cavity. 

a. Lower margin of fang-groove with 4 teeth m-inatur. 

I). Lower margin of fang-groove with 3 teeth. 
«!. Central posterior eyes one diameter apart ; apical 
portion of embolus shorter, more distinctly 
flanged on both sides ; 3rd tooth more remote ; 

tibial spur less broad and flattened loiir/i/arsis. 

hK Central posterior ej-es less than half a diameter 
apart ; apical portio!i of embolus longer, less 
conspicuously Hanged on the lower margin ; 3rd 
tooth not remote ; tibial spur broader and more 
flattened e.rfensa . 

B. Embolus of bulb without any erect black spur in its 

basal cavity. 
a-. Tibial spur straight, blunt at apex. 

rt'*. Tibial spur forming a cons])icuoas angle on its 
inner maririn. Tooth 3 remote from tlie others; 
apex of embolus less broadly flanged on its upper 
margin maccauncUi. 

/;■'. Tibial spur without angle on inner margin. 
Teetli equidistant; apex of eml)olus more broadly 

flanged on its upper margin connexa. 

h'-. Tibial spur strongly curved, ]iointed at apex amazoniea. 


A. Lower mavgin of fang-groove witli 5 teeth, the Ith 

very small, situated on the inner side between the 
3rd and otli. Vulva without any central tongue-like 
sclerite, but with two small blunt cusps in tlie middle 
of the anterior margin, and an obliq\ie elongate 
lateral lobe on each side, converging behind tlie 
cusps kpj/.ierli>i(ji. t<p. ii. 

B. Lower margin] c)f fang-groove with 3 (or 4) teeth. 

Vulva with a well-defined central tongue-like sclerite. 
a. Lower margin of fang-groove with 4 teeth, the 3ril 

smaller itriiui/ur E. Hhnuu. 


h. Lower margin of fang-groove with 3 subeqnal teeth. 

«'. Size much larger, 25 mm. Leg iv. at least ten 

times longer than the carapace. Third tooth on 

lower margin of the fang-groove slightly more 

remote from the second than the latter is from 

the first. 

a". Central tongue-like sclerite much narrower, 

dumb-bell-shaped, about eqtiallj' dilate at each 

end ellacomhei, sp. n. 

J-. Central tongue-like sclerite much broader, 
almost as broad in the middle as posteriorly, 
much broader anteriorly but somewhat vari 

able longitarsis C. L. Koch. 

h^. Size much smaller, 11 mm. Leg iv. not more 
than 5-6 times longer than the carapace. Teeth 
on lower margin of fang-groove equidistant. 
Central cavity of vulva long and narrow, straight 
on the inner margins ; central tongue elongate, 
not dilate anteriorly, but narrowed amasonica, sp. n. 

1. Trechalea longitarsis (C. L. Koch). (Plate XV. figs. 13-17.) 

Die Arach. xv. p. 65, t. 522. fig. 1462 (sub Triclaria). 

? Trechalea longitarsis E. Simon, Ann. Soc. Ent. Belg. xlii. 
1898, p. 20. 

Type $ , in coll. Mus. Imper. Berlin. Hah. Colombia. 

Although the identity of this species cannot be satisfactorily 
settled without reference to the tyjje-speci'men, if it still exists, 
I give a desci'iption and figures of the characters of a form with 
annulated legs which was taken in Colombia. If the type no 
longer exists, and no more exact locality than " Colombia " can 
be ascertained for the type- specimen, then the examples here 
described stand as ^'- topoti/pes" — examples from the same locality 
in which the type was taken — and will serve as a standard of 
comparison for the species in the place of the original type. 

Had tSimon given more definite characters (for the majority of 
the species of Trechalea possess 3 teeth) which would enable us to 
recognize his identification of T. longitarsis with any certainty, 
and if these (for he makes no mention of the locality) were taken 
in Colombia, then his examjdes would be the recognized " topo- 
types " to which we should have to refer for an identification of 
T. longitarsis (C. L. Koch). 

Present identification of T. longitarsis (sec. F. P.-Cambr.). 

Total length, S 20, ? 25 mm.— $ . Leg i. 61 ; ii. 68 ; iii. 55 ; 
iv. 75 mm. Tib. i. 15 ; iv. 16"5 mm. Prot. iv. 21 mm. 

Examples in coll. Brit. Mus. from Colombia. 

(J 5 . — Colour. Carapace rich brown-black, with a pale orange 
/V-shaped spot behind the eyes, having a central dusky line, and 
one on each side ; a pale rufous-white, narrow, submarginal 
band, and a narrow black velvety marginal band. Ahdomeyi 
brown, with a black central foliated band : a more exact description 
is impossible from dried and wrinkled examples. Sternum and 
ventral area dull yellow-brown. Legs brown, very distinctly 
annulated with rufous-white hairs, protarsi less distinctly, tarsi 


not at all. Mandibles black, clothed with hoaiy-white hairs in 

Structure, General characters similar to those of other species of 
the genus. Central posterior eyes one diameter apart (lass in S ) ; 
clypeus six times as high as the diameter- of an anterior central 
eye (in J four times). 

Vulval area broader than long, rounded, rectangular ; central 
tongue distally broad, widely dilate basally ; variable in diliereut 

Palpus. Tarsus twice i\s long as broad ; tibia two-thii-ds the 
length of the fcxrsus ; one-fourth longer than the patella. Tibial 
spin- broad basally, curved apically, terminating in a small hook, 
having on the inner side a slightly serrated ridge. The embolus 
of the bulb has a stout erect spur springing from the cavity below 
the apex of the lamina of the bulb, while the embolus itself is 
short, strongly curved, and conspicuously flanged on each side. 

Ilah. CoLOMBJA, dried examples ( c? »& $ ) ; Venezuela, dried 
examples ( d" tfc $ ) ; also many examples in spirit, in coll. Brit. 

2. Trechalea urinator E. Simon. (Plate XV. figs. 3-5 a.) 
Ann. Soc. Ent. Belg. xlii. p. 20 (1898). 

Type S , gynetype $ , in coll. E. Simon. Total length ( c? , $ ) 
20-25 mm.— (c?)- Carap. 11. Leg i. 74; ii. 82; iii. 62; iv. 
85 mm. Tib. i. & iv. 19-5 mm. Prot. iv. 22 mm. 

Both sexes with 4 teeth on the lower margin of the fang- 
groove. Legs 4, 2, 1, 3. Clypeus 3-3-1- times in height the 
diameter of an anterior central eye. 

Carapace, abdomen, and legs entirely without dark bands or 
annules, or these are at least very faint ; entii'ely clothed A\'ith 
dusty olive-bi'own hairs and pubescence. 

Palpus. Tibia 1| times longer than broad ; tibial spur slender, 
its apex bent over, forming a single small hook. The inner 
apophysis deeply impressed across the middle, forming two blunt 

Vulva with a short broad central tongue, broadly dilate basally. 

If I am correct in my identification of Simon's species, it is a 
common Spider tlii'oughout Ecuador. 

Hah. Ecuador : Guayaquil, Loja ; Rio Durango ; Bulun ; 
Cachavi ; Cavondelet ; Paramba ; Salidero {Rosenhery). 

3. Trechalea ellacombei, sp. nov. (Plate XV. fig. 6.) 
Type $, in coll. Brit. Mus. Total length 25 mm. Cai-ap, 

10 mm. Leg i. 68; ii. 75; iii. 60; iv. 80 mm. Tib. i. 17; iv. 
17-5 mm. Prot. iv. 22-5 mm. 

Lower margin of fang-gi-oove with 3 teeth. Legs 4, 2, 1 , 3. 
Clypeus 4 times the height of the diameter of an anterior central 
eye. The whole of the central area of the carapnce l)lack, with 
margins and stria? flavous. Abdomen too much shrivelled to admit 
of the coloration being recorded. Legs very deeply streaked mid 

Proc. Zool. Soc— 1903, Vol. I. No. XI. 11 


annulated with black, except the tarsi. Yulva with a long centi'al 
tongue, dilate basally, and its apex incurved. 

This species is nearly allied to T. i(,rinator, but may be recognized 
by the annulated legs, the possession of only 3 teeth on the 
mandible, and by the form of the vulva. 

Hah. Surinam, Bergen Daal (collected by Mr. C. W. Ellacombe). 

4. Trechalea macconnelli Pocock. (Plate XV. figs. 7, 8.) 
Trans. Linn. Soc. Lond., Zool. (2) viii. 2, p. 67, Sept. 1900. 
Type c? , in coll. Brit. Mus. Total length 16 mm. Carap. 

9 mm. Leg i. 64; ii. 70; iii. 55"5 ; iv. 73. Tib. i, & iv. 17 mm. 
Prot. iv. 18 mm. 

Lower margin of fang-groove with 3 teeth. Legs 4, 2, 1, 3. 
Clypeus twice the height of the diameter of an anterior central 
eye. Carapace with a large black patch in the centre, its margins 
scalloped by the flavous margin round the carapace and the pale 
strife. Abdomen dusky with a flavous pattern and two flavous 
spots above the spinners. Palpus : Tibia not, or scarcely, longer 
than broad. Tibial spur slender, concave lamelliform, its lower 
edge slightly serrate ; convex and incurved at the apex ; seen from 
above it has a A-shaped emargination. The inner apophysis is 
cylindrical, low and impressed across the top. 

Sab. British Guiana : Mt. Roraima (collected by Messrs. 
Quelch & McConnell). 

5. Trechalea connexa (0. P.-Cambr.). (Plate XV. figs. 9, 10.) 
Triclaria connexa 0. P.-Oambr. Biol. Centr.-Amer., Arach. 

Aran. i. p. 233, t. 30. figs. l,la-e{S ), 1898. 

Trechalea connexa F. P.-Cambr. loc. cit. ii. p. 313, t. 30. fig. 17 
{6), 1902. 

Type S , in coll. Godman & Salvin. Total length 14 mm. 
Carap. 7 mm. Leg i. 40 ; ii. 45 ; iii. 39 ; iv. 48 mm. Tib. i. 10; 
ii. 11 mm. Prot. iv. 13"5 mm. 

Lower margin of fang-groove with 3 teeth. Legs 4, 2, 1, 3. 
Clypeus as high as two diameters of an anterior central eye. 
Carapace flavous, streaked and mottled with brown. Abdomen 
dark brown. Legs decidedly annulate. Palpus : Tibia half longer 
than broad ; spur, seen from below, without any hook at its apex. 
Embolus of palpus without two ridges as in T. 'macconnelli, shorter 
and more broadly flanged on its anterior margin. 

Ilab. Mexico, Atoyac {R. H. Smith). 

6. Trechalea EXTENSA (O. P.-Cambr.). (Plate XV. figs. 11, 12.) 
Triclaria extensa 0. P.-Cambr. loc. cit. i. pp. 174, 233, t. 22. 

figs. 10, I0a-f{s), 1896. 

Trechalea extensa F. P.-Cambr. loc. cit. ii. p. 313, t. 30. figs. 16, 
16«,6(c?), 1902. 

Type S , in coll. Godman & Salvin. Total length 20 mm. 
Carap. 10 mm. Leg i. 69 ; ii. 77 ; iii. 58 ; iv. 82 mm. Tib. i. 
17; iv. 19 mm. Prot. iv. 20 mm. 


Lower margin of fang-groove with 3 teeth. Legs 4, 2, 1, 3. 
Clypens 3 times as high as the diameter of an anterior centiul 
eye. Carapace unicolorous flavous, margined with dusky black. 
Abdomen unicolorous brown, with a short central anterior pale 
dorsal bar. Legs faintly annulate. Palpus : Tibia twice as long 
as broad ; spur, seen from above, stout, projecting, cnrved, with 
a stout sharp cusp or hook on the inner side at the apex, very 
similar to but much stouter than in T. imnator. Inner apophysis 
deeply impressed in the middle. Embolus as in T. long liars is , 
but less bi'oadly flanged. 

ITab. Guatemala {>Sarg). 

7. Trechalea amazonica, sp. nov. (Plate XY. figs. 18-20.) 

Type d", gynetype ? , in coll. Brit. Mus. Total length ^ 10; 
( 2 15 mni.). Carap. 6 mm. Leg i. 31 ; ii. 32 ; iii. 29 ; iv. 37*5 
mm. Tib. i. &, iv. 7-8 mm. Prot. i. & v. 10 mm. 

Lower margin of fang-gi'oove with 3 teeth. Clypeus in height 
equal to l^j diameters of an anterior central eye. Legs 4, 2, 1, 3. 
Carapace dull stx'aw-3'ellow ; margins and sti'i;^ slightly suffused 
with dusky brown, produced by fine dai'k hairs ; mottled with 
patches of white hairs, angular on mai'gins ; clypeus fringed and 
clothed with hoaiy- white hairs. Mandiljles, pedipalps, and a spot 
on ocular area behind the eyes clothed with hoary- white haii's. 
Abdomen olive-green mottled with brown and hoary- white. 
Ventral area clothed with silver- white hairs. Sternum and legs 
dull yellow beneath, the latter brighter above ; femora indistinctly 
annulated with dusky brown above ; base of patella black. Tibite 
and pi'otai'si with two broad dark annulations. Pedipalps also 
annulated with brown. 

Vvira with a narrow oblong central cavity, its lateral margins 
straight and parallel, with an elongate central tongue, not dilated 
at either end, not incurved at apes, without any black shining 
bosses on each side. 

The male and female here described wei-e taken from the same 
trunk of a tree on a swampy island in the largo opposite tSautarem. 
They were ci-ouching on the bark lilce Spai'assitls, and when dis- 
turbed moved with great rapidity. 

Hah. Lower Amazons : Santarem ; Breves, Parana Buyassu ; 
Pard {F. P.-C. 1895-6). 

8. Trechalea keyserlingi, sp. nov. (Plate XY. figs. 1, 2.) 

Type $, in coll. Keys. Brit. Mus. Total length 20 mm. 
Carapace 9'75 mm. Leg i. 44; ii. 46; iii. 36; iv. 49 mm. 
Tib. i. 11 ; ii- 11 "5 mm. Prot. iv. 13 mm. 

Lower mai-gin of fang-groove with 5 teeth. Clypeus five times 
as high as the diametei' of an anterior centi'al eye. Legs 4, 2, 
1, 3. Carapace with two broad centi'al bands, widely separate 
behind the eyes, connivent behind ; tlie clear space l)ehind Hie 
eyes marked with lateral and central fine streaks. Abdomen with 
a broad deep-brown foliate band above, consti'icted just behind 



the middle and again slightly above the spinners, broadly margined 
with pure white pubescence. Femora of legs deeply blotched and 
streaked above with black ; tibise distinctly, protarsi indistinctly 
annulated. Vtdva without any central tongue, but with two 
small cusps in the middle and two separate posteriorly converging 
lateral sclerites, quite distinct in form from that of any other 
species here recorded (see Plate XV. fig. 2). 

This specimen M^as identified by Keyserling as T. longitarsis 
C. L. Koch. See p. 157. 

Hah. Brazil, Rio Grande do Sul {Keys. Coll.). 

Genus Hesydrus Simon, 1898 (June 30). 

Simon, Hist. Nat. Ar. 2, ii. p. 315. Type, H.'pahistris Simon. 


The species belonging to this genus are very closely allied to 
Trechalea, but foim a small group chiefly distingu^ishable by the 
glabrous, gibbous mandibles, with a well-marked carina on the 
outer side, in the male sex. The legs in both sexes are subequal 
in pairs 4-2, 1-3 ; being also much shorter in proportion than in 
Trechalea. Both margins of the fang-groove with 3 teeth. Eyes 
as in Trechalea^ central posteriors slightly over one diameter 
apart ; clypeus low and porrected. Tibia and protarsus i. with 
2-2-2-2 spines beneath; protarsi 1-4 with a small central apical 
spine beneath. 

In Biol. Oentr.-Amer., Arach. Aran. ii. p. 305, I have referred 
the species given as the tyjoe of Hesydrus (sec. 2 a, H. jidlieni) 
to Enna O. P.-Cambr., and the name Hesydrus is ajDplicable to the 
species under sec. \a with H. jyahistris as the type. It appears 
that Simon has somehow confused the characters of these two 
groups. In Enna (including H.jibllieni) undoubtedly the clypeus 
is moi-e vertical ; the centi-al posterior eyes are two diameters 
apart, or moi'e, and the anterior row is not wider than the 
posterior central row ; while the tarsi are straight and not flexible. 
But in H. palustris (for if not identical, the form before me 
is certainly congeneric with this species) the clypeus is pori'ect, 
the central posterior eyes one diameter apart, the antei'ior row 
wider than the posterior central, and the tarsi flexible ; the eyes 
of the posterior row are certainly not equidistant, a character 
given for distinguishing Hesydrus from Trechalea. Fortunately, 
however, the citation of a definite type species enables one to 
rectify the confusion. 

The two species known to me may be recognized by the 
following charactei's : — 

a. Tibial spur of palpus, seen from above its apex, not bifid, 

slightly emarginate and prominent at its angles, but 

not incurving in the form of two teeth habilis 0. P.-Cambr. 

b. Tibial spur of palpus, seen from above its apex, deeply 

bifid, its angles forming two sharp teeth curving in- 
wards and downwards palustris Simon. 


1. Hesydrus talustris Simon. (Plate XV. figs. 22-25.) 
Ann. Soc. Ent. Belg. xlii. 1898, p. 21 ( J , $ ) ^ 

Type d'jgynetype $, in coll. E. Simon. Total length cj" 11 
($16 mm.). — Caiap. 6 nnn. Leg i. 25; ii. 30; iii. 24-5; 
iv. 30 mm. ' Tib, i. 6 ; iv. 7 mm. Prot. iv. 10-5 mm. 

Lower margin of fang- groove with 3 teeth. Legs 4-2, 1-3. 
Clypeus porrect, as high only as the diameter of iin anterior 
fu>ntral eye. Cai'apace dull flavoiis with dusky haii's. Falpas : 
Til)ial spur short, concave lamelliforni, deeply bifid at its apex, 
forming two small, sharp, incurving teeth. Inner apophysis elon- 
gate, not impressed across the middle, produced at its untci-ior 
apical angle to foiiu a small cusp. Vulva with a broad central 
cavity, more than two-thirds the transverse width, concave on its 
lateral margins, having i\. long nai-row central tongue, not dilate 
at either end, with a shiny black boss on each side, on the lloor 
of the cavity ; very vai'iable in foi-m. 

Hah. Ecuador: Vta-Ainh-A^Roseuhery). Loja, Zamora\ — Vene- 
zuela : Merida. 

It is not certain that my identification of this species is correct, 
but it is highly probable. 

2. Hesydrus habilis (0. P.-Oambr.). (Plate XV. fig. 21.) 
Tridaria habilis O. P.-Cambr. loc. cit. i. pp. 173, 233 t 22 

figs. 9, 9«-/(c?), 1896. 

Trechalea habilis F. P.-Cambr. loc. cit. ii. p. 313, t. 30 fig 15 
(c?), 1902. 

Type S in coll. Godman and Salvin. Total length 10 mm. Carap. 
5 nun. Tib. i. 5-5 mm. (other legs broken off or mutilated). 

Lower margin of fang-groove with 3 teeth. Logs 2, 4, 1, 3 
(sec. O. P.-C). Clypeus as high as 1| diameter of "an anterior 
central eye. Legs dull yellow-brown, slightly annulate. The 
colours of the cai^apace and abdomen appear to have faded. 
Mandibles glabrous shiny and very gibbous dorsally. Palpus : 
Tibia one-third longer than broad. Ti])ial spur, seen from above 
its apex, not bifid, its angles not forming sharp incurving teeth. 

Hah. Guatemala: Costa Rica (/S'cw*/, iioyers). 

3. Hesydrus estabanensis E. Simon. 

Ann. Soc. Ent. Belg. xlii. 1898, p. 20 ( J , ? ). 

Type S , gynetype $ , in coll. E. Simon. Total length" J $ , 7 mm. 

One would susjiect, from the descriptions, that this species is 
congenei'ic with //. palusiris, the males having the luaudililes 
convex and cai-inate on their outer apical margin. 

Hah. Venezuela : San Esteban. 

4. Hesydrus bucculentus E. Simon. 
Ann. Soc. Ent. Belg. xlii. 1898, p. 20 ( c5' ). 
Type S , in coll. E. Simon. Total length 10 mm. 

The remarks made under the last species apply also to this. 
Hab. Brazil : Rio {Gounelle). 


Fam. SenoculiDjE. 
Genus Senoculus Taczanowski. 

1872. jSenoGiohcs Tacz. Hor. Soc, Ent. Ross. ix. p. 106. — Type, 

aS'. maronicids. 

1873. Lahdams 0. P.-Cambr. Proc. ZooL Soc. Loncl. p. 118.— 

Type, L. monastoides. 
1880. Platyctenihs Keys. Yerh. z.-b. Ges. Wien, sxix. p. 338, 
note. Nom. nov. for Senocidus (nom. inappro. sec. Keys.). 
1880. Stenoctemis Keys. Yerh. z.-b. Ges. Wien, xxix. p. 340, — 

Type, S. gracilis. 
1883. Neothereutes Holmb. Bol. Acad. Nac. Cienc. Cordoba, v. 

p. 35. — Type, N. darwini Holmb. 
In a former paper, " Cteniform Spiders from the Lower Amazons 
&c.," Ann. Mag. Nat. Hist. (6) xix. p. 90, Jan. 1897, 1 have tabu- 
lated the characters of the known species of this genus so far as 
I could diuw reliable characters from the existing descriptions. 

The table is here re-modelled and includes additional species 
from Central America, piiblished in the ' Biologia Centrali- 
Amei'icana,' and a new species occurring in the British Museum 
Collection, of which a diagnosis is given below. I have not seen 
S. 2)ci,rallehts Simon, but it is certain that my example from the 
Lower Amazons belongs to that species, since M. Simon has kindly 
compared a figure of the vulva with that of his own type. 

The following is a list of the species already published : — 

1872. S. maronicus Tacz,, 5 , St. Laurent de Maroni. Hor. Soc. 

Ent. Ross. ix. p. 106, t. iii. fig. 4. 

1873. S. monastoides (0. P.-Oambr.), 5 , Brazil, Rio Grande. 

P.Z. S. Lond. p. 118. 

1879. S . riohromacidatus Keys., $, Peru. Yerh. z.-b. Ges. 

Wien, p. 339, pi. iv. fig. 30. 

1880. S. pl'umosus (E. Sim.), $ , Brazil, Para. Bull. Soc. Zool. 

Fr. p. 154. 
1880. >S'. pttrjnireus (E. Sim.), $ , Panama. Bull. Soc. Zool. Fr. 

p. 155. 
1880. S. paraUelus (E. Sim.), $ , Brazil, Tefife. Bull. Soc. Zool. 

Fr.p. 156. 
1880. S . riificapillus (E. Sim.), c? , Brazil, Para. Bull. Soc. 

Zool. Fr. p. 154. 
1880. .S'. iricolor (E. Sim.), S $ , Brazil, Tefi'e. Bull. Soc. Zool. 

Fr. p. 153. 
1880. S. gracilis (Keys.), $ , Peru, Aimable Maria. Yerh. z.-b. 

Ges. Wien, p. 341, t. iv. fig. 29. 
1883. S. dartoini (Holmb.), $, Formosa, Chaco, Arg. Rep. 

Bol. Acad. Nac. Cienc. Cordoba, v. p. 35. 

1896. S.2^rolatus (0. P.-Cambr.), S 2, Mexico, Atoyac. Biol. 

Centr.-Amer., Arach. Aran. i. p. 218,t. 28. figs. 3, 3a-f{ $ ). 

1897. tS. pallidus (F. P.-Cambr.), S , Brazil, Rio Janeii-o. Ann. 

Mag. Nat. Hist. (6) xix. p. 92, pi. iv. fig. 5 (sub 



1902. S. canal icalatus F. P.-Cambr., J $ , Paiiaiiia, Biigalia. 
Biol. Oentr.-Amer., Aracli. Aiun. ii. p. 350, t. 33. 
figs. 3(c^), 4, 4«($). 
The characters by which the species may be distinguished, so 
far as I can gather them from the descriptions, ai-e as I'dHows : — 


A. Carapace and abdomen clothed with plumose hairs, iricolor Siiiioii. 

B. Carai)ace and al)domeu clothed witli simple hairs. 

I. Tibia i. and ii. with 9 — 10, protarsi with 8 — 9 long 

siiiiius beneath on each side j)alUdus F. I'.-C'auilir. 

II. Tibia i. and ii. with 4 — i long spines beneath, pro- 

tarsi with o — 5. 

a. Anterior portion of ocular area almost vortical . ruJlcapiUus Siuion. 

b. Anterior portion of ocular area porrcet, liori- 


1. Apex of lamina round tlio inner anierinr por- 

tion of the bulb of the jmlpus nmeli broader 

and more deeply and broadly bifurculato ... in'olatus (_). I'.-Cainbr. 

2. Aj)ex of lamina round the inner ant('ri<ir 

portion of the bulb of tlie palpus much 
narrower and less deeply bifurculate (simply 
bitid and canaliculate) canalicidatus F. l'.-(.'ambr. 


A. Carapace and abdomen clothed witli hairs. 

1. Carapace deeply striate, central stria oval-elon- 

gate; tarsi i. and ii. with 4 — i, protarsi i. and ii. 

with 3 — 3 long spines beneath ■plamosiis Simon. 

2. Carapace scarcely striate, central stria short, sub- 

punctiform; tibia i. and ii. with 5 — o, protarsi 

i. and ii. with 4 — 4 long spines beneath iricolor Simon. 

B. Carapace and abdomen clothed with simple hairs. 

a. Legs very long ; tibia i. and ii. with long spines 

beneath, 10 on inner side, 9 on outer side. 
«'. The lateral lolies of the vulva bearing a rounded 

boss on the outer margin just behind the 

middle (see Keyserling's tigiire) ffracilis Keyserling. 

b^. The lateral lobes of the vulva without any 

rounded boss parallchis Simon. 

b. Legs sliorter, tibia i. and ii. with only 4 — 1- or 

5 — 5 long spines beneath. 

aa. Tibia i. and ii. with 5 — 5 long spines beneath. 

(i'. Posterior eyes very large, centrals less than a 

diameter ajiart, scarcely Vt, diameters IVom 

til'' laterals; anterior portion of ocular area 

almost vertical ruJicajiilJns Simon. 

b-. Posterior eyes moderate in size, centrals over 
a diameter apart, at least two diameters 
from the laterals. Anterior jjoi'tion of 

ocular area jjorrect luonaaloidi s (). I'.-t'ainbr. 

bh. Tibia' i. and ii. with 4. — '1 long si)iues beneath. 
«•'. Protarsi i. and ii. with 3 — 3 long spines 

beneatli, besides lateral and apical s|)ines. . (dhiilKn V. l'.-('anil)r. 

i''. Protarsi i. and ii. with 6— o long spines 

beneath, besides lateral and ajiieal sjiines. 

a'K External sclerites of vulva evenly rounded 

on their outer nuirgin, sinuously concave 

on their innei margin, their ai)ices 

(anterior pcn-tion) sliarj), with a mimite 

cusp on their outer margin prolalus O. P.-Cand)r. 

b*. External sclerites of vulva emarginate on 
their outer sides, evenly convex on their 
inner margins, their ajiices forming a 
broad hook, not very sharp at the point, 
without anyminute cusp canaliviilahiit F. P.-Cambr. 


Senoculus albidus, sp. nov. (Plate XIY. fig. 14.) 

T3rpe 2 > ill coU. Brit. Mus. Total length 14 mm. 

In general appearance this species is similar to others of the 
genus, the legs being clothed with a lateral fringe of curving 
simple silky white hairs. The abdomen is also clothed at the 
apex on each side with long pure- white hairs. The vulva consists 
of two large oblong convex sclerites convergent forwards, and 
somewhat concave at this point, see Plate XIY. fig. 14, where 
also the vulva of *S', parallelus Simon is figured (fig. 13). 

Hah. Brazil ; Rio Janeiro. 

Note. — OiS. purpureiis (E. Simon) and 8. darwini (Holmberg) 
I cannot give any reliable characters. S. inaronicus Tacz., the 
type of the genus, is, according to the author, immatvire, and no 
mention is made of the number of spines on the legs. 

Plate XIV. 

Fig. 1. TJiaumasia velox, $ , vulva, p. 154. 

2. „ annulipes, $ , vulva, p. 154. 

3. Dossenus marginatus, $ , p. 155. 

4. „ „ $ , vnlva. 

5. „ „ ? , eyes. 

6. Paradossenms nigricans, $ , p. 155. 

7. „ „ ¥, vulva. 

8. „ „ ? , eyes. 

9. „ J) (? 3 tibial spur of palpus. 

10. Thanatidius spinipes, ? , eyes, p. 156. 

11. „ „ ditto, in front. 

12. „ „ ? , vvilva. 

13. Senoculus parallelus, $ , vulva, p. 166. 

14. „ albidus, $ , vulva, p. 168. 

Plate XV. 

Fig. 1. TrecJialea Iceyserlingi, ^ , p- 163. 

2. „ „ vulva. 

3. „ MWMaior, ^5 tibial spur of palpus, p. 161. 

4. „ „ ?, vulva. 

5. „ „ $ , embolus of palpus. 

5 a. „ „ (? , „ from below. 

6. „ ellacomhei, ? , vulva, p. 161. 

7. „ maccowMeZZi, (j, tibial spur of palpus, p. 162. 

8. „ ,, (J, embolus of palpus. 

9. „ connexa, $ , tibial spur of palpus, p. 162. 

10. „ ); (? ) embolus of palpus. 

11. „ ea-^emsa, <J , tibial spur of palpus, p. 162. 

12. „ „ (J , embolus of palpus. 

13. „ longitarsis, $ , embolus of palpus, p. 160. 

14. „ „ (? , ditto, apex enlarged. 

15. „ „ (J, tibial spur of palpus. 

16. „ „ (J , ditto, example from Venezuela. 

17. „ „ ? , vulva, example from Colombia. 

18. „ amazonica, <? , tibia of palpus and spur, p. 163. 

19. „ „ $ , embolus of palpus. 

20. „ „ ? , vulva. 

21. Hesydrus Jiahilis, $ , tibia of palpus and spur, p. 165. 

22. „ jpalustris, $, tibia of palpus and spur, p. 165. 

23. „ „ c? J embolus of palpus. 

24. „ „ $ , riglit mandible, from outer side. 

25. „ „ ?, vulva. 

P Z.S. 1903, vol.1 PI .XVI. 

,, S.6.9 do. PICTUS 

12 P.Z.S.1903.vol.I.P).XVJI 


■ Th.C 

of ppcL - 

TTh or pp 

C.C del. 


iSale &:DamelssoiiI.\'^ litK. 



2. On the Marine Fanna o£ Zanzibar and British East Africa, 
from Collections made by Cyril Crossland in the Years 
1901 and 1902.— Polych^ta. Part I. By Cyril 
Crossland, B.A., B.Sc.^ 

[Received Jaimaiy 15, 1903.] 
(Plates XYI. & XVII.^) 
The collections referred to were made under the following 
circumstances : — Sir Charles Eliot, K.C.M.G., C.B., late Fellow 
of All Souls College, Oxford, at present H.M. Consul-General at 
Zanzibar and Commissioner for British East Africa, took me out 
with him as his private assistant in his researches on Nudibranch 
Mollusca. I made collections not only of this, bvit of the other 
marine groups ; accounts of which, by various specialists, will 
appear in these 'Proceedings' from time to time. The largest 
collections are those of the Nudiln-anchiate Mollusca and Polychrete 
Ann-elids, of which groups about 150 species in each are to be 
described. Su^ Charles Eliot has ah^eady publi.shed one part of the 
results of his examination of the Nudibi-anchs (P. Z. S. 1902, vol. ii. 
p. 62) ; other papers by various authors wih shortly be ready. 

For the benefit of other possible woi'kers in this region, I may 
mention that the greater part of my collections was made in two 
localities, viz. Chuaka Bay, on the east coast of Zanzibar, and 
Wasin Harbour, near the Anglo- German boundary on the main- 
land. ^ The former locality is extremely rich in shore forms, and 
dredging in 3 fathoms of water at the north side of the mouth of 
the bay was often very productive. Wasin Harbour averages a 
depth of 10 fathoms, and here I collected almost entirely by 
dredging. The bottom is extensively covered by a species of 
Telesto, the branches of which are overgi'own by an encrusting red 
sponge. Among this a great variety of the smaller Polycha?ta 
and Nudibi-anchiata are found. 

My mention of these two localities is of the more importance to 
future workei's because large stretches of the coast are extremely 
barren. Unfortunately this applies to the two principal towns of 
East Africa, and especially to Mombasa. A few miles from 
Zanzibar, e. g. round the islands and sandbanks which surround 
the harbour, and on one portion of the shore about a mile to the 
south of the town, at low spring-tide only, are lich collecting- 
grounds, and near this latter area dredging is profitable, especially 
at a depth of 5 fathoms. The greatest portion of the Zanzibar 
channel is, however, extremely barren. 

More detailed descriptions of the reefs of East Africa, with 
maps, will be found in my two papers : " On the Coral Reefs of 
Zanzibar," and " Pemba and British East Africa," in the Proc. 
Camb. Phil. Soc. vols. xi. & xii. (1902). 

1 Conimnnicated by Prof. W. C. JIcIntosh, F.Z.S. 

2 For expLinatiou of the Plates, sec p. 176. 


The gratitude of all zoologists is thus due to Sii^ Chai'les Eliot, 
whose generosity and scientific zeal have enabled these collections, 
the first of considerable size from this region, to be made. I 
wish also to exjoress my best thanks to Mr. Stanley Gardiner, who 
helped me by his advice and in every other possible way, both 
during my residence in East Africa and in the working out of the 
collections on my return to England. 



The position of the genus Phyllochcetoptents among the lower 
Chpetopteridfe is shown by the following table of the genera : — • 

A. Notopodia of the second body-region not foliaceous 

{i. e. two body-regions onl^') Ranzania. 

B. With foliaceous notopodia posteriorly. 

1. Body divided into two regions Telepsatus. 

2. Body divided into three regions. 

C One pair of tentacles Spioch^topterus. 

<.Two pairs of tentacles Phtlloch^itoptekus. 

The most characteristic feature of the Chsetopteridse is the 
adaptation of certain parapodia for the production of a respiratory 
current, which modification, completed in the genus Chcetopterus, 
makes the latter one of the most remarkable forms of animal life. 

This peculiarity is not developed in Ranzania, whose only 
Chsetopterid features are its general build of body, which is 
divided into an anterior flattened muscular and glandular portion 
bearing long notopodial setse only, and a rounder weaker posterior 
portion with delicate notopodia and neuropodia of uncinigeral 
tori ; in addition it is tubicolous in its mode of life, and procures 
food by the ciliated grooves of its tentacles and dorsal surface. 

To these features are added notopodial gills in Telepsavus, in 
which case they are developed on every segment of the hind-body. 
In the remaining two genera this modification is restricted to 
more or fewer of the middle segments, to two only in Spiochceto- 
pterus and several species of PIiTjllochcetopterus, but up to 25 in 
other species of the latter. 

In all the eight species of PJiylloclicetopterus yet known the 
body is vermiform, the notopodial gills comparatively small and 
simple, bifid, and containing capillary setse. All the setpe are 
characteristically Chsetopterid, in the first region long with leaf- 
like ends, and arranged in a row as in Chcetopterus, though the 
notopodia of P. acictoUgerios alone approach in their long pointed 
shape those of the former genus. In the fourth notopodium of 
the first body-region one or more setee are enlarged and of a dark 
brown colour, but are not flattened as in Chcetopterits. The uncini 
are of one kind only throiighout the body, but their form is more 
specialized than in Chcetopterus, and is that which occurs in, for 
example, the Sabellidse. The parapodia ax-e alike in all the sj)ecies, 
differing only in the proportions of the parts, excepting the hind- 
body notopodia of P. aciculigerus (6). 


The prostomium is, in this family, usually more or less reduced. 
In Chcetopteriis itself it is completely merged with the oral 
funnel, and the same condition is described for some Phyllocha-to- 
pterids (e. g. P. claparedi IVIcIntosh). In all the members of this 
genus it is normally scarcely distinguishable externally, both pairs 
of tentacles being carried by the peiistomium. In certain specimens 
of F. elioii, however, as described below, it is quite prominent, 
and so was discoverable in all the four species I liave examined. 

Tlie first body-region is an almost solid mass of muscular and 
glandular tissue in all species, but jiosteriorly musculature is 
confined to the ventral surface, except at the level of the parapodia, 
where muscle-bands surround the body. Between these the body- 
wall is excessively thin doi-sally and laterally, and always black 
fi'om the colour of the gut-walls. 

Up to the present six species have been described, viz. : — ■ 

P. socialis Clapar^de. Mediterranean and Atlantic (1) (4). 

P. majoi" Clapar^de. Mediteiranetin (1). 

P. gracilis Langerhans. Canary Islands (2). 

P. claparedi Mcintosh. Japan (3). 

P. aciculigems Crossland. The IMaldive Islands (6). 

P. gardineri Crossland. The Maldive Islands (6). 

To these I now add : — 
P. elioti. Zanzibar. 
P. piictus. British East Africa. 

Clapar^de's P. fallax was founded on such chai'actei'S as the 
numbei'S of segments composing the three body-i-egions, the 
i-inging of the tube, &c. ; these have been shown by Roule (4) to 
be variable in members of this genus. 

The principal distinctions of systematic importance are : — 

(1) Presence or absence of eye-spots, and of the development 

of the prostomium, 

(2) The number of the strong setfe in the notopodium of the 

fourth setigerous segment. 
(.3) The proportionate sizes of the paiis of the parapodia of 

the regions B and C 
(4) Ringing and other such details of the tubes are variai)le, 

but (1) in some species they are sti-aight and occiu" 

singly, and (2) in others they are of a U -shape and occur 

in numbers twisted together. 

These characters are distributed as follows : — 

(1) Eye-spots present in P. socialis, P. major, P. elioti, P. 

gardineri, P.pictus; absent in P. claparedi, P. acicidigerus, 
P. gracilis. 

(2) A single strong seta in 4th notopodiiun in P. socialis \ 

P. major, P. claparedi, P. pietus. More than one seta 
strengthened. P. gracilis with three, P. gardineri three, 
P. elioti two, and P. acicidigerus eight. 

1 In P. sncialis two setie may exceptionally occur, hut the extra one is alwaj's 
smaller than the one it accompanies. 


(3) The parapodia of the region consist of double neuro- 

podial tori and a clavate papilliform notopodium containing 
a single seta in all the species except P. aciculigerus and 
P. claparedi. 

(4) ^ Tubes straight, occurring singly in P. claparedi, P. elioti, 

P. onajor. Twisted together, in numbers, P. socialis and 
P. pictus. 

Phylloch/etopterus elioti", sp. n. (Plate XYI. figs. 1-3 & 8 ; 
Plate XVII. figs. 10-13.) 

Of this species numerous specimens occur in Chuaka Bay, 
Zanzibar. The sti-aight, stiff, opaque black tube, 6 to 9 inches 
long, is buried in the sand at low tide, one or two inches only 
projecting. Its presence is usually rendered conspicuous by the 
growth of a tuft of bright green enteromorpha on the projecting 

The colour of the living animal is milk-white anteriorl}^, and 
black posteriorly. There is no pigmentation, and the brilliant 
blue and yellow colours described by Clapai'ede and Lo Bianco in 
the Naples species are not found in either of the two species I 
have seen alive. The black colour of the gut gives the vTSiial 
green solution in alcohol. 

The peristomium is very mobile, the whole shape of the head 
being thus very different in difierent specimens. When expanded, 
its shape is as in PI. XYI. fig. 1 . In a few cases only, where the 
head is much contracted and bent back, does the prostomium and 
its eye-spots come into view as in fig. 2. 

The first pair of tentacles are very slender and long, attaining 
a length of 9 mm. 

The fore-body measures, in a large specimen, 6 mm. in length 
by 2'5 mm. in breadth, and consists of about 14 setigerous 
segments. (There were 13 in three specimens, 14 in three more, 
and 15, 16, and 17 each in three more. The average is thus 14 
in nine specimens.) The notopodia are very short and stumpy, 
but the setfe and their arrangement, as in the other species of the 
genus, recall Chcetopterus. The fourth foot has two strong setfe 
on each side, rarely three on one side or the other, though in one 
specimen there were three on one side and four on the other. 
The ordinary setse are all straight. 

The region B (fig. 3) consists of from 20 to 25 segments, the 
parapodia of which are very like those of P. gardineri, except that 
the notopodial flap is slightly smaller, and there is a long space 
between it and the ventral portion of the neuropodium, and the 
parts of this latter are of more nearly equal size. The neuropodium 
of the first segment of the gill-region is not divided. There are in 
each notopodium five long setse whose flexible heads project. The 
uncini (fig. 8, c) are triangular with very fine teeth, just visible 
under a ^-inch objective. 

The hind-body has more than 25 segments, but none of my 

1 The tubes of Gardiner's two species were uiifortuiiatelj^ not obtained. 

2 Thus named in honour of Sir Charles Eliot, K.C.M.G., C.B., H.M. Consul- 
General at Zanzibar, and Commissioner for British East Africa. 


specimens are quite complete. The notopodia are the usual 
clavate papilla? containing a single seta. The neuropodia ai-e as 
in the gill-be<aring region, but much smaller. 

The two species P. elioti and P. ga7-dineri are closely related, 
differing only in the reduction of the peristomial collar in the 
latter and the proportions of the mid-body parapodia and sette. 

The only work on the intei-nal anatomy of a member of this 
genus is the section of the fore-body of P. claparedi given by 
Mcintosh, although examination of tlie anatomy of tlie species 
of Pliylloclufiiopterus would be extremely interesting, both for 
comparison among themselves and with the other geneiu of the 
family \ especially Chcetoj)terus, Having neither time nor full 
opportunity for this woi-k, to which my inclinations ai-e most 
strongly drawn, I can only give hei'e a few deductions from the 
examination of a series of sections of a specimen of this genus, 
which appear to differ in many minor details from the above- 
mentioned section of P. claparedi. 

Body-ioall. — The region A is, as shown by the sections figured 
(PI. XVII. figs. 11 & 12), an almost solid mass of glandular 
and muscular tissue, in marked conti'ast to the delicacy of the 
i-egions B and C. The whole of the ventral epithelium is sti'ongly 
glandular beyond the limits of the well-mai-ked ventral shield, and 
these glands extend anterioi'ly to the dorsal surface and even on to 
the prostomium. The cuticle is of extreme delicacy, if really 
present. The musculature consists of weak circular and very 
strong longitudinal muscles, but the division of the latter into 
two dorsal and two ventral bundles cannot be made out. In the 
regions B and C the only muscle occurinng doi'sally is a very 
delicate circular layer (PI. XVII. fig. 13). Diagonal fibres can be 
ti'aced in the region A between the nei've-cords and the notopodia, 
which become more definite muscles in the region B (fig. 13). 

The nervous system is in contact with the skin. The two 
ventral cords lie at a distance from one another in the region A, 
but approach one another posteriorly (cf. figs. 12 & 13), their 
arrangement corresponding thus to that in Cluciopterus. Trans- 
verse commissures, portions of one of which are shown in PI. XVII. 
fig. 12, connect the two cords. 

The brain (PI. XVII. fig. 10), which in Chcetopterus is a narrow 
band dorsal to the mouth, not differentiated from the circum- 
cesophageal commissures, is here more distinct, in correspondence 
with the presence of the prostomium in this genus and its absence 
in the case of Choitojiterus. It is a perfectly simple swelling of the 
circum-oeso]ihageal commissures. The continuity of the nervous 
system with the epidermis is shown by a comparison of the 
sections of the prostomium given in figs. 10 & 11. The former 
shows the circumoeso2)hageal connectives, that on the left as it 
crosses from the venti'al to the dorsal side, that on the right, near 
the eye-spot (e) passing into the prostomium. The z-eplacement 
of the ordinary glandular epithelium of this area by the densely 

* For anatomy of Telepsavus, sec pi. xiii. and its explanation in Claparede's 
' Annelides ISedeutaires.' 


staining nerve- nuclei of the brain (which are similar to those 
seen on the outer side of the ventral nerve-cords &c.) is shown 
distinctly, this replacement being complete in fig. 10, where the 
epidermis is composed entirely of nerve- cells. 

The eye- spots are a pair of groups of cells, each containing 
numerous minute granules of black- brown pigment. These are in 
continuity with the nerve-cells of the biuin, but are anterior and 
dorsal to its fibrous part. 

The coelom is small and broken up into several distinct portions. 
The largest of these are a pair of spaces lying laterally and 
ventrally to the gut (fig. 12), bounded dorsally by the powerful 
muscles of the seta-sac. Dorsally and medianly, commencing as a 
space in the prostomium, is a third pai't which posteriorly becomes 
a mere canal surrounding the doi'sal blood-vessel. In the region B 
the coelom is more normal, thoiigh reduced in size. It remains 
divided into right and left halves by the dorsal and ventral 
mesenteries of the gut. 

The vascular system consists of dorsal and ventral vessels, both 
running in the gut-mesenteries. The former breaks up at the 
base of the prostomium into three branches (fig. 10), and large 
connecting-vessels are found in the anterior segments. Posteriorly 
the dorsal vessel forms a laig-e sinus covering the dorsal wall of 
the gut (fig. 13). 

Alimentary canal. — The mouth is richly ciliated, and the outer 
ends of its columnar epithelial cells contain a few minute specks 
of the black-green pigment so characteristic of the family. These 
are absent from the narrow triangular gut of the region A, but 
reappear in great quantities in the larger thick- walled alimentary 
canal of the regions B and C (fig. 13). 

The gut-lining here consists of long crowded cells, the swollen 
distal ends of which are ci-owded with minute specks of the 
pigment, indicated by dark dots in the figui'es. 

The transverse sections of the notopodial gills, shown in fig. 13, 
are interesting. The space which they contain is ccelomic, and at 
its centre is a'bundle of five or six setae wrapped closely together 
by fleshy tissue. The transverse section of this resembles that of 
a telegraph-cable, the wires being represented by the set*. At 
each end of the oval section is seen a sharply-cut groove, the 
sides of which are of granular, rather deeply- staining protoplasm, 
without nuclei, supporting externally very long cilia. Laterally 
is a more deeply-stained area of the epithelium, perhaps of nervous 

Phylloch^topterus picttjs, sp. n. (Plate XYI. fig. 5.) 
The tubes of this species are brown and translucent, quite free 
from sand or mud. They were found clustered together in con- 
siderable numbers on the under side of a large stone at low-water 
level on Pungutiayu islet, Wasin. Each tube has two openings, 
each limb of the U being about 4*5 cm. long. The bending is 
irregular, and they are twisted and fused together, so that it is 
usually almost impossible to separate out any one tvibe. 


The contained worms are the smallest of the species descrilied 
here, or in (6), being slightly smaller everyway than in P. elioti. 

The specific name refei'S to the abundance of bi-own colour 
found on their tentacles and fore-body. Pigmentation, except of 
the gut, is rare in the ChK>topterida\ Of the other species, it is 
descril)ed in only two, viz. F. clcqxiredi (3) and P. socicdis (1) : in 
both these it is comparatively slightly developed. 

The ground-colour of tlie anterior region, tentacles, and para- 
podia of P. jnchis is, in life, creamy white. The greater part of 
the posterioi- regions is, as usual, black, by reason' of the pigmen- 
tation of the gut. 

On either side of the groove which runs along tlie upper surface 
of tlie larger tentacles are regularly arranged squarish blotches 
of brown, as shown in PI. XVI. fig. 5. Brown dots are scattered 
also over the pro- aiid peristomium and the anterior segments of 
the fore-body. There are two dark bands with indefinite edo-es 
along the ventral bases of the notopodia, and a broad crescent 
across the ventral surface of the end of the region A. 

The movith is not at all funnel-shaped, but rather slit-like, tlie 
two small peristomial lappets coming together from either side. 
The prostomium is large, flattened from side to side, and projects 
above the dorsal surface of the fore-body. The very slender second 
setigerous pair of tentacles lie close on either side of it, being thus 
very inconspicuous. Two elongated, but distinct, eye-spot?, of a 
dark brown colour, occur one on each side of the prostomium. 

The legions A and B consist of the following numbers of 
segments in difl^erent individuals : — 

A ... 15 16 — 13 15 12 15 1*3 

B... 7 9 5 7 5 3 8 — 

* These specimens were below the average size. 

The fourth notopodium has but one large seta of the shape 
shown in PI. XVI. fig. 9, a. The bending of this seta is remarkable 
as occurrnig only in this species. Its three teeth, when seen from 
one point of view, giwe an explanation of Langerhans's figure of 
the corresponding seta in P. gracilis, which is not otherwise 

The other setae are all straight, and present no peculiarities 
except those of the last notopodium, which are bent, and the head 
is finely toothed (fig. 9, c). 

The parapodia of the region B (figs. 5 & 6) are small, the 
neuropodial ridges especially so. In correspondence with this the 
uncini are very minute and delicate (-03 mm. long), and their 
teeth barely visible under a |-inch objective (fig. 9, d). The 
neuropodia do not extend far up the side of the body, and the 
space between them and the notopodial gills is not filled up by 
the triangular membranous gill-flap as in the other species (fig. 6). 
The notopodia contain two or three thin seta? which do not project." 

In the region the notopodia are reduced to little clavate 
papilla?, containing one seta, as in all the other species except 



[Feb. 17, 

P. daparedi and P. acicidigerus (see fig. 6, of P. elioti). The 
neuropodia also are small. 

List of the Literature. 

(1) Claparede. — Annelides du Golfe de Naples, 1868. 

(2) Langerhans. — " Ueber einige Canarischen Anneliden." Nov. 

Act. der Leop.-Carol. Deutschen Akad. Bd. xliii., 1881. 

(3) McIntosh.—' Challenger ' Reports, vol. xii., 1885, p. 374. 

(4) Roule. — " Oampagne du Caudan." Ann. d. I'Universite de 

Lyon, 1896. 

(5) Lo Bianco. — Ann. Tub. nel Golfo di Napoli, 1893. 

(6) Crossland. — J. Stanley Gardiner's ' Fauna and Geography 

of the Maldive Archipelago.' In the press. 


Fig. 1. Normal condition of the head of Plii/llochcetopterus elioti (p. 172), as seen 
from dorsal side. X 10. 

2. Side view of a specimen whose prostomium and eye-spots are visible. X 10. 

3. Side view of three segments of the region B. X 10. 

4. Ventral view of the same. 

5. P. pictus (p. 174). Dorsal view of anterior part of hodj\ X 10. 

6. Side view of three segments of the region B. X 10. 

7. A notopodium of the region C in either species. X 100. 

8. Setffi of P. elioti (p. 172). 

(a) Thickened seta of fourth parapodium. X 83. 
(6) Normal seta from third foot. X 90. 
(c) An uncinus. X 1000. 

9. Seta3 of P. picttis (p. 172). 

(a) Thickened seta from the fourth foot. X 100. 

(h) Head of a normal seta from the same bunch. X 200. 

(c) From last foot of region A. The shaft is bent and part of the head 

finely toothed. X 100. 
{d) One of the uncini. X 1000. 

Plate XVII. 

Fig. 10. Section of Fhyllochcetopterus elioti (p. 172) through the brain, bases of 
short tentacles, &c. 

11. Section a little anterior to fig. 10, through mouth, first feet, &c. 

12. Section at level of fifth foot. 

13. Section near beginning of region B. All sections X 42. 

The meaning of the lettering on the figures of the Plates is as follows :— 

cJrc.TO.=circular muscle. 
COS. =ccBlum. 
(^.6».= dorsal blood-vessel, 
(^.ct?.= dorsal ciliated groove. 
c^.cc6.=dorsal division of ccelom. 
«:Lwettr.= dorsal division of the neuro- 
^iZL=gill-flap between noto- and 
^L= glandular areas or epithelia 

Im. = longitudinal muscle-layer. 
5w.= muscle. 
m. qfppd.=mMscles of parapodium. 

«•.= nerve. 
w.c.=nerve-cells. = nerve-cord. 
»»/■.= nerve-fibres, 
wejo/j. =nephridium. 
weMr.=neuropodium or ventral division 

of the foot. 
wo^o.= notopodium or dorsal division 

of the foot, 
joeri. =peristomium. 
^ji3f^.= parapodium or foot, 
jsro. =prostomium. 
ti &io = first and second pairs of ten- 
■y.6«.= ventral blood-vessel. 
•i).MeM>',= ventral division of the neuro- 

p. Z . S. 1903, ATolIPLXVIII. 

M.P.T>a.rker ]ifk. 





--- h-dt: 


- cck. o.p.- 


Pourkeir & "West inap. 



3. Oil the Axis, Athis, ami Froatliis in the Higher 
Theriodonts. By R. Bkoom, M.D., B.Sc, C.M.Z.S. 

[Rocoived Jaiuuiry 20, 1903] 

(Plate XVIII.") 

Hitherto, tliouyli something has been known of tlie structure 
of the axis in at least two of the Dicynodont.s and in (J>/iiO(jiiathus, 
we have been practically ignorant of the structure of tlu; atlas in 
any of the Theriodonts or Anoniodonts. In fact, Prof. iSeeley (1) 
in descrilnng the axis of CyaoynatJius, seems to be in doubt 
whethei- he is really dealing with an axis, or with an axis and 
atlas cond)ined. On page 100 of his paper he says, "The first 
vertebra appears to be anch}dosed to tlie second " ; while a little 
further on he states that " this vertebi'a is remarkable for its 
form being exactly like the odontoid pi'ocess of the vertebra in 
many animals, and suggesting the idea that the atlas is lost " ; 
aud in a note he adds : '' This appears to be confii-med by the sub- 
setjuently to be described condition in Tropidostomci duiini." In 
counting the vertebrse, however, he counts the axis as the 1st 
cervical, and states that "there are 6 cervicals." Gadow (2), in 
his recently published work on Reptiles, possibly misled by Seeley, 
definitely states that " the atlas is fused Avith the axis." Con- 
sidering how mammaldike the higher Theriodonts are, and that 
the axis bears a very marked resemblance to the mammalian axis, 
having a large spine and a well-developed odontoid process, it 
seems remarkable that the idea should have arisen that the atlas 
was anchylosed to the axis, more especially as there is no part 
of the axis in Cijnognathios that bears any resemblance to any 
known atlas. 

When in Grahamstown recently, I had an oppoitunity, through 
the kindness of Dr. Schonland, of examini)ig the Theriodonts in 
the Albany Museum, and especially the very fine specimens of 
Gomjjhognathus and Trirachoclon^ which have been described by 
Seeley (3), and which have been so marvellously developed under 
his direction. In both the type specimens of Gomphognathus 
kannemeyeri and Trirachodon kannemeyeri the anterioi' cervical 
vertebrae are most beautifully shown, but Seeley in his desciiption 
of the specimens makes no i-eference to this most important 
region, beyond stating (p. 53) that " the occipital plate [in Trira- 
cliodoii\ is not completely exposed owing to some of the vertebrae 
being in contact." Two of his figures of the skull of Trirachodon 
show the atlas and part of the axis ; but as no distinction is shown 
between inati-ix and bone, it is ilifiicult to make anything of the 

The beautiful specimen which forms the type of (Joinphogiudhns 

' For explanation of the Plate, sco ji. 180. 

Proc. Zool. Sue— 1S)03, Vol. I. Xo. Xll. 12 

178 DR. E. BROOM ON THE [Feb. 17, 

consists of two complete mandibles in articulation with the 
posterior half of the skull. The lower jaw lies almost at right 
angles to the axis of the skull, and the upper cervical vertebrae 
have been to some extent protected by passing down between the 
two rami. The skull consists of the practically complete occiput 
with both tempoial arches in almost perfect condition, and with 
most of the parietal and sphenoidal regions well shown. Though 
the sutures unfortunately are not distinct, the occiput is so well 
preserved, practically without distortion, that I think it worthy 
of being figured. 

The vertebrae so far as preserved consist of the proatlas, atlas, 
axis, and the 3rd and 4th cervical vei'tebrae. 

The atlas {at.) consists of a well-developed arch and a distinct 
hypapophysis { The arch is partly broken in the middle, but 
must have borne considerable resemblance to that of the mammals. 
It differs in having had a distinct posterior zygapophysis { 
for articulation with the small anterior zygapophysis of the axis 
{ The zygapophysis of the left side is well presei-ved, but 
that of the right has been broken off. On the right side the 
outer part of the arch is continued backwards as a well-developed 
bony process or rib { It is probable that the process is 
really a rib, but I have been unable to find a suture between it 
and the arch. On the under surface, what appears to be part of 
the rib of the left side is seen lying by the side of the body of the 
axis { The well-developed hypapophysis is also well seen. 
It appears to have had only a ligamentous connection with the 
arch of the atlas. 

In front of the arch of the atlas, and lying in the hollows formed 
above and outside of the occipital condyles, are the two portions 
of the proatlas. Each consists of a short bony arch with a 
well-developed j)rocess passing upward and outwai'd fi'om its 
outer end. ^ 

The axis is well developed, and bears considerable resemblance to 
that of Cynognathus. The spine forms a very large flattened crest. 
In the specimen it is somewhat damaged, but there is probably 
not a great deal missing. In front, the spine passes forwards 
over the arch of the atlas, and possibly forms an articulation 
with it as in the Monotremes. Between the anterior part of the 
spine and the odontoid process is the small anteiior zygapophysis 
{ The odontoid process (o.j».) is well developed, and on the 
right side it is seen articulating with the arch of the atlas. On 
the under surface the body of the axis is seen, as also a part of the 
odontoid process in articulation with the hypophysis of the atlas. 
On the upper surface, near the posterior end of the centrum, on 
the right side is a portion of the rib of the axis. 

The 3rd and 4th cervicals present no features worthy of any 
special remark. Remains of ribs are seen in connection with 

In the type specimen of Trirachodon Jcannemeyeri there are 
preserved and well shown the two portions of the proatlas, the 


arch and liypapophysis of the atlas, and the odontoid process and 
part of the body of the axis. Tliough the structure of the 
vei'tebrse is essentially similar to that in Gomphogiiathus there are 
many points of diSerence. 

The proatlas {2).a.), though occupying the same relative position 
as in Gomphoy)iat}tas, is less specialized. It is present as a pair of 
curv^ed, moderately thin, bony plates lying in fi'ont of, and i^robably 
somewhat overlapping, the ai-ch of the atlas. There is no bony 
process developed as in Uomi)liognatlius. 

The atlas («^) consists of a well-developed arch and a distinct 
hypapophysis. The arch is not so complete as in Gompliof] iiaihas, 
anil must have differed considerably in regai'd to the airangement 
of the antei'ior articular surfaces. In Gompliognathiis the occi- 
pital condyles {p.c!) are small and moderately close together, 
whereas in Trirachodon the condyles {p.c)) are veiy lai-ge and as 
wide as the atlas. The arch, perhaps on this account, does not 
close in inferiorly to meet the hypapophysis, with which it can 
only have been attached by ligament. Tiie zygapophysis {^ 
for the axis is very small. The hypajjophysis { is very 
similar to that in Gomphognatlius. 

Of the axis only the odontoid process, imperfectly displayed, 
and a part of the body remain. 

The structure of the upper cervical vertebrae is so imperfectly 
known in the majority of fossil I'eptiles, that there is some diffi- 
culty in satisfactorily dealing with the affinities of the structures. 
It is difficult to avoid being struck by the close resemblance of 
the axis, atlas, and proatlas in the Theiiodont to those structures 
in the Crocodiles, more especially as there is no close affinity 
between the groups. The explanation is probably to be found in 
the fact that both the Crocodiles and tlie Theriodonts, though far 
removed from each other, have retained with but slight modifi- 
cation the type of structures met with in their common ancestor. 
In Hphenodon, though the proatlas is small, the tj^pe is practically 
the same ; and I have recently discovered that Procolophon has a 
well-developed proatlas. Frocolophon has usually hitherto been 
associated with the Dicynodonts, Theriodonts, and Pareiasaurians, 
but as it has a persistent notochord, abdominal ribs, a plate-like 
pubis and ischium, and 2, 3, 4, 5, and 4 phalanges in the digits, it 
is manifestly much more closely lelated to PalcnohatUrla than it is 
to the Tlieriodonts and Dicynodonts. In most of its structures it is 
probably as primitive as the common ancestor of the Crocodiles 
and Theriodonts, so that there is good reason for believing that 
the common ancestor had a well- developed proatlas and pi-obably 
an atlas and axis veiy similar to those found in later forms, 
but almost certainly with the odontoid process as a distinct 
element, as seen in Ichthyosaurus. Ko proatlas has yet been 
detected in Pareiasaurus, but I have found one in Lystrosaurus 
{^ = Ptychognaihas Owen). It will probably yet be found in mo«t 
of the primitive reptilian types. 


180 MR. C. TATE REGAN ON THE [Feb. 17, 

Wo7'ks referredj to. 

(1) H. G. Seeley : " On the Skeleton in new Cynodontia from 

the Karroo Rocks." Phil. Trans, vol. 186 B. p. 59 (1896). 

(2) H. Gadow. ' Amphibia and Reptiles.' London, 1901. 

(3) H. G. Seeley : '' On the Gomphodontia." Phil. Trans, vol. 186 B. 

p. 1 (1896). 


Fig. 1. Occiput and upper cervical vertebras of G-ompliognathus Tcannemeyeri. Nat. 

Fig. 2. Under view of atlas and axis of G-ompliognatlnts. Nat. size. 
Pig. 3. Posterior view of atlas &c. of Trirachoclon Icannemei/eri. Nat. size. 
Fig. 4. Under „ „ „ „ ,, Nat. size. 

Fig. 5. Side „ „ „ „ „ Nat. size. 

at., atlas ; ax., axis ; e.o., exoccipital ;, lij'papophysis of atlas ; o.c, occipital 
condyle; o.p., odontoid process ; jp.a., proatlas ; r**, r'*, ribs ;, atlas rib;, 
axis rib; sq., squamosal;, zygapox^liysis of atlas;, zygapoplij'sis of axis; 
3c, 4o, 3rd and 4th cervical vertebrae. 

4. A Eevision of the Fishes of the Genus Triacanthus. 
By C. Tate Regan, B.A.^ 

[Received January 20, 1903.] 

Although six species of Triacanthus were described and figured 
by Bleeker in the ' Atlas Ichthyologique,' GUnther, in his Catalogue, 
recognized only three (viz. : Tr. strigilifer Cantor, biacideatus 
Bloch, and hrevirostris Schlegel), and did not accejat either of the 
four described by Dr. Bleeker as new, but placed three of them 
(viz. : Tr. macriirus, hlochi., and oxycephalus) in the synonymy of 
Tr. hiaculeatus^ and the fourth (^Tr. nieuhofi) in that of Tr. hrevi- 
rostris. Since then this arrangement has not been challenged, nor 
has any new species of this genus been described. 

Subsequent to the reading of my paper on the Plectognathi ^, I 
examined the specimens of Triacanthi in the British Museiun 
Collection, which include Bleeker's types, and I have come to 
the conclusion that all six species described by Bleeker are valid, 
although the one he called Tr. niacnvrus is certainly identical with 
Tr. hiaculeatus Bl., a species not recognized by him ; whilst a 
seventh species, which has been generally confused with Tr. hrevi- 
rostris Schleg., is now described for the first time as Tr. indicus. 
A complete revision of the synonymy has thus become necessary, 
and the need for more complete diagnoses of the various species is 
obvious. In the descriptions given below, which are in each case 
based on several specimens, the total length is measured to the 
base of the first caudal ray, the length of head to the gill- opening, 
the length of the caudal peduncle from the base of the last dorsal 
to that of the first caudal ray, the length of the snout from its tip 
to the vertical from the anterior margin of the eye, that of the 

1 Communicated by G. A. Boulengee, F.R.S., V.P.Z.S. 
" P. Z. S. 1902, ii. p. 284. 



postor1)ital part of the head from the vei-ticnl from tlie posterior 
margin of the eye to the gill-opening. Young specimens have a 
shorter snout, a deeper body, and longer dorsal and ventral spines 
than adults, and have been excluded fi-om the diagnoses. In each 
case the total length of the largest example here described is given. 

Triacaxthus Cuvier. 

Body compressed, covei'ed with small rough scales ; caudal 
peduncle more or less elongate, tapering ; mouth small ; teeth in 
2 sei'ies in each jaw, 10 in each outer sei'ies, incisor-like ; those of 
the inner series obtuse, rounded, 6 in the upper j:iw, 2 in the 
lower. Branchiostegals 6 ; pseudobranchite present. Spinous 
dorsal with 5 rays, the first a long and strong spine ; soft dorsal 
with 20-25 rays; anal with 15-20 rays; ventral fins each repre- 
seiited by a strong spine, without soft rays ; caudal widely forked. 
Vertebrae 20. 

In all the species the colour is strikingly similar, being bluish 
grey above, silver)^ below. The only tangible difierences in colour 
are that the membrane of the spinous dorsal fin is in some specimens 
immaculate, in otheis wholly or pai-tly blackish. 

Key to the Species. 

I. Second raj' of spinous dorsal more than half as 

long as tlie first, membrane of spinous dorsal 
immaculate ; length of base of anal half that of 
soft dorsal; 1). V, 20-22 ; A. 15-16 1. T>: sln<jilife)-CM\ior^ 

II. Second ray of spinous dorsal less than half as 

long as the first. 

A. Snout concave ; pelvis between ventral spines 

tapering to a distinct point posteriorly. 
a. jMcnibranc of spinous dorsal immaculate; 

length of caudal peduncle 4}r-5| times iu 

total length ; length of base of anal l^-lf 

times in that of" soft dorsal ; D.V, 22-23; 

A. 16-17 2. Tr. hJoahi Blocker. 

h. Membrane between first two rays of spinous 

dorsal blackish ; length of caudal peduncle 

4.f-5 times in total length; length of liase 

of anal Ij-l^ times in that of soft dorsal; 

D.V, 23-24; A. 18-19 3- Tr. biaculcatus Bloch. 

c. Membrane of spinous dorsal immaculate; 

length of caudal peduncle 63-7 times in 

total length; length of base of aiml Ij!- 

IJ times in that of soft dorsal; D. Y, 

21-25; A. 17-19 4. Tr. TAcckcr.. 

B. Snout straight ; pelvis between ventral spines 

sairceU" narrowed posteriorly; D.V, 24-25; 
A. 18-20. 
a. Membrane of spinous dorsal blackish ; snout 

half as long as the head ; postorbital part of 

head not shorter than cye-diametf^r 5. T>: breviroslris ^chlogel. 

h. Membrane of spinous dorsal blackish ; snout 

more than halt as long as the head ; post- 
orbital part of head shorter than eye-diameter. 6. Ti: iHiiifKS, n. sp. 
c. Membrane between first two rays of spinous 

dorsal blackish ; snout less than half as long 

as the head ; postor6ital part of head shorter 

than eye-diameter 7. Tr. nieuhofi Sleeker. 

182 MR. C. TATE REGAN ON THE [Feb. 17, 

Triacanthus STRIGILIPER. 

Triacanthus strigilifer Cantor, Mai. Fish. p. 363, pi. ix. (1847); 
Bleeker, Atlas Ichtli. v. p. 89, pi. ccxxix. fig. 3 (1865); Giintliei", 
Cat. viii. p. 211 (1870). 

Triacantlmis longirostris Hollard, Ann. Sci. Nat. (4) i. 1854, 
p. 46, pi. ii. fig. 3. 

Depth of body about equal to the length of head, abovit 3 
times in the total length, length of caudal peduncle 4|-5 times. 
Snout very slightly concave, its length 1|—1| times in that of the 
head, eye-diameter 3-4 times, interorbital width 4-5 times. 
Interorbital space concave, without distinct median ridge. Upper 
edge of occipital ci'est in the same straight line as that of the 
snout ; distance to base of first dorsal spine from posterior margin 
of eye about 1^ times the eye-diameter. D. V, 20-22 ; A. 15-16 ; 
the first dorsal spine longer than the head, the second considerably 
more than llalf as long as the first, the others much shorter ; the 
base of the anal fin about half as long as that of the soft dorsal 
fin ; pelvis between the ventral spines moderately broad, tapering 
to a point posteriorly. Membrane of spinous doi'sal immaculate. 

Total length 185 mm. 

Hah. (Seas of Arabia and India ; East Indian Archipelago. 

Triacanthus blochi. 

Triacanthus hlochi Bleeker, Nat. Tijds. Ned. Ind. iii. 1852, 
p. 81 ; Atlas Ichth. v. p. 89, pi. ccxvii. fig. 1 (1865). 

Triacanthus hiaculeatus (part.) Giinther, Cat. viii. p. 210 

Depth of body about 3 times in the total length, length of head 
3|-3|- times, length of caudal peduncle 4^-5^ times. Snout 
concave, its length 1|— 2 times in the length of head, eye-diameter 
3-3| times, interoi'bital width 4-5 times. Interorbital space 
concave, with median ridge scarcely, if at all, distinct. Upper 
edge of occipital crest forming an angle of about 160° with that 
of the snout; distance from posterior margin of eye to base of 
first doi'sal spine about 1| times as long as the eye- diameter. 
D. Y, 22-23 ; A. 16-17; the first dorsal spine longer than the 
head, the others short; length of base of anal fin 1|-11 times in 
that of the base of soft dorsal fin. Pelvis between ventral spines 
narrow, tapeiing to a slender point. Membixme of spinous 
dorsal immaculate. 

Total length 150 mm. 

llah. East Indian Ai'chipelago ;_ China. 

Triacanthus eiaculeatus. 

Balistes hiaculeatus Bloch, Ausl. Fische, pi. 148. fig. 2 (1785). 
Triacanthus hiaculeatus Cuv. Begne An. ii. p. 152 (1817). 
Triacanthus angustifrons HoUard, Ann. Sci, Nat. (4) i. 1854, 
p. 45, pi. ii. fig. 2. 


Triacanthus macrwus Bleeker, Atlas Ichth. v. p. 91, pi. cexxii. 
fig. 3 (1865). 

Triacanthus hiaculeatus (part.) Giinther, Oat. viii. p. 210 (1870). 

Depth of body about 24 times in the total length, length of 
head about 3| times, length of caudal peduncle 4|-5 times. 
Snout concave, its length 1^14 times in the length of head, eye- 
diameter 3jT-4 times, intei'oi-bital width 4-5 times. Intei'oi'lntal 
space concave, with median lidge scarcely, if at all, distinct. 
Upper margin of occipital crest forming an angle of about 170° 
with that of the snout ; distance fi'om postei-ior mai-gin of eye to 
base of first dorsal spine lf-l|- times as long as the eye-diameter. 
D. V, 23-24; A. 18-19; the first dorsal spine longer than the 
head, the others short ; length of base of anal fin If-li times in 
that of the base of the soft dorsal fin ; pelvis between vential 
spines rather narrow, tapeiing posteriorly to a slender point. 
Alembrane between first and second rays of spinous doi'sal fin 

Total length 180 mm. 

Hah. East Indian Archipelago ; Australia ; China. 

Triacanthus oxycephalus. 

Triacanthus oxycephalus Bleeker, Yerh. Bat. Gen. xxiv. 1852, 
p. 27, pi. v. fig. 10 ; Atlas Ichth. v. p. 90, pi. ccxx. fig. 3 (1865). 

Triacanthus hiaculeatus {'^•Avt.)(3i\x\\t\\eY, Cat. viii. p. 210 (1870). 

Depth of body 2y— 2| times in total length, length of head 
about 3 times, length of caudal peduncle 6^—7 times. Snout 
slightly concave, its length about 1-i times in the length of head, 
eye-diameter 3-4 times, interorbital width 3-3j times. Inter- 
orbital space flat. Upper edge of occipital crest convex ; distance 
from postei'ior margin of orbit to base of first dorsal spine about 
1| times as long as the eye-diameter. D. V, 24-25 ; A. 17-19 ; 
first dorsal spine longer than the head, the others short ; length of 
base of anal fin 14-1 1 times in that of the base of the soft dorsal 
fin ; pelvis between the ventral spines broad anteriorly, tapeiing to 
a point posteriorly. Membrane of spinous dorsal fin immaculate. 

Total length 140 mm. 

Hah. East Indian Archipelago. 

Triacanthus brevirostris. 

Triacanthus hrevirostris Schlegel, Faun. Japon., Poiss. p. 294, 
pi. cxxix. fig. 2 (1846); Bleeker, Atlas Ichth. p. 94, pi. ccxxxi. 
fig. 3 (1865); Gunther, Cat. viii. p. 209, part. (1870). 

Triacanthus hiaculeatus Bleekei', Yerh. Bat. Gen. xxii. 1849, p. 6. 

Triacanthus rhodopterus Bleeker, Yerh. Bat. Gen. xxiv. 1852, 
p. 25, pi. iv. fig. 8. 

Triacanthus russellii Bleeker, t. c. p. 25. 

Depth of body 21 -2| times in the total length, length of head 
3^-31 times, length of caudal peduncle 41-54 times. Snout 
straight, its length about twice in that of the head, eye-diameter 
3|-5 tijEces, intei-orbital width about '6^ times. Length of post- 


orbital part of head equal to or greater than eye-diameter. Inter- 
orbital space with a more or less distinct median ridge, with a 
groove on each side of it. Occipital crest convex, moderately 
elevated, becoming nearly horizontal in front of the base of the 
first dorsal spine. D. V, 24-25; A. 18-20; first dorsal spine 
shorter than the head, the others short ; length of base of anal fin 
1|— 1|- times in that of soft dorsal fin ; pelvis between ventral 
spines broad, scarcely narrowed posteriorly. Membrane of spinous 
dorsal fin blackish. 

Total length 250 mm. 

Hob. East Indian Archipelago, Seas of China and Japan. 

Triacanthus indicus, n. sp. 

Russell, Indian Fishes, p. 14, pi. xxi. (1803). 

Balistes hiacideatus Bennett, Fishes of Ceylon, pi. xv. (1830). 

Triaccmihus hiaculeatus Cantor, Mai. Fish. p. 360 (1847); Day, 
Fishes of Malabar, p. 260 (1865). 

Tricicanthus hrevirostris HoUard, Ann. Sci. ISTat. (4) i. 1854, p. 45, 
pi. ii. fiff. 1 ; Giinther, Cat. viii. p. 209, part. (1870) ; Day, Fishes 
of India, p. 685, pi. clxxv. fig. 1 (1878). 

Depth of body 2^2 1 times in the total length, length of head 
3|-3| times, length of caudal peduncle 4|-5 times. Snout 
straight, its length about 1| times in that of the head, eye-diameter 
3|-4| times, interorbital width 3-3| times. Length of post- 
orbital part of head distinctly less than eye-diameter. Interorbital 
space with a more or less distinct median I'idge with a gi'oove on 
each side of it. Occipital crest elevated, its upper edge nearly in 
the same straight line with that of the snout. D. Y, 24-25 ; 
A. 18-20 ; first dorsal spine nearly as long as the head, the others 
short ; length of base of anal fin 1^-1 1 times in that of soft dorsal 
fin ; pelvis between ventral spines bi'oad, scarcely nai-rowed 
posterioi'ly. Membrane of spinous dorsal fin blackish. 

Total length 220 mm. 

Hah. Coasts of India from Kurrachee to Penang ; Ceylon ; 
Andaman Is. 

This species difiers from T. hrevirostris chiefly in the longer 
snout, the shortei' postorbital pait of head, and the shoi-ter and 
more elevated occipital crest. The figures given by Russell and 
Bennett are excellent. 

Triacanthus nieuhofi. 

■^ Nieuhof, Gedenkw. Zee en lantr. p. 272, fig. 

Willoughby, Ichthyology, Appendix, p. 5, pi. x. fig. 2 (1686). 

? Gronow, Mus. i. p. 52 (1754-6) & ZoophyL p. 53 (1763-81). 

Triaca7ithics 7iietchofi Uleeker, Verb. Bat. Gen. xxiv. 1852, p. 26, 
pi. iv. fig. 9, & Atlas Ichth. v. p. 92, pi. ccxvii. fig. 3 (1865). 

Triacanthus hrachysoma Bleeker, Nat. Tijds. Ned. Ind. iv. 
1853, p. 128. 

1 I have not hcew nble to verily tliis reference. 


"i Balistes hipes Gronow, Cat. Fish. p. 37 (1854). 

Triacanthushrevirostris (part.) Giinther, Cat. viii. p. 209 (1870), 

Depth of body about 2^ times in the total length, length of 
head 3g-3|- times, length of caudal peduncle ^l^— 4^ times. Snout 
straight, about 24- times in the length of head ; eye-diameter about 
3j times and equal to interorbital Avidth. liOngth of postorbital 
part of head less than eye-diameter. Inteioi'bital space with 
median ridge with a groove on each side of it ; occipital crest 
strongly elevated, the distance fi'om the base of fii'st dorsal spine 
to the upper angle of gill-opening rather more than the distance 
fi-om the base of the fii'st dorsal spine to the anteiior margin of 
eye. D. V, 24-25 ; A. 18-20; first dorsal spine longer than the 
head, the others shoit ; length of base of anal fin about 1 h times 
in that of the base of soft doi-sal fin ; pelvis between the venti'al 
spines broad, not much narrowed posteiiorly ; membi*ane between 
first two rays of spinous dorsal fin blackish. 

Total length 126 mm. 

Hah. East Indian Archipelago. 

This species has a shorter and more declivous snout than 
Tr. hrevirostris, and also occipital crest more elevated, postoi'bital 
part of head shorter and body deeper. 

5. On the Geographical Variations o£ the Sand-Yiper, Vipera 
amuwdytes. By G. A. BouLENGER, F.ll.S., V.F.Z.S. 

[Received January 30, 1903.] 
(Text-figures 27 & 28.) 

The variations of Vijyei-a ammodi/ies in connection with the 
distribution of the sjiecies have not I'eceived sxifiicient attention. 
Having succeeded in bringing together and carefully compaiing a 
series of 55 specimens from various localities, I have convinced 
myself that the South-eastern specimens (Gi'eece, Archipelago, 
Syria) can be distinguished from the typical form from Austi-ia- 
Hungaiy, Dalmatia., Bosnia, and Montenegi'o, not by means of 
any single absolute character, but by a combination of characters, 
as shown by the following definitions : — 

Forma tijpica (text-fig. 27 a). — Naso-rosti'al shield usually ^ 
reaching the canthus rostralis, and extending considerably higher 
up than the upper border of the I'ostral, which is usually broader 
than deep (text-fig. 28 a) ; rostral " horn " with 3 (rai'ely 2 or 4) 
transverse seiies of scales between the rostral shield and the 
apex. Ventral shields 145 to 163. The dark shade on the lower 
lip, if present, broken up by liglit bars sepai'ated by 2 to 4 labial 
shields. Lower surface of end of tail usually red ^. Grows to 
80 centimetres. 

Var. meridlonaUs (text-fig. 27 b). — ISTaso-rostral shield never 

1 5 exceptions out of 30 specimens examined. 

" Yellow in one specimen from the Dinaiic Alps', Bosnia. 



[Feb. l\ 

reaching the canthns rostralis, and hut rarely extending higher 
tip than the upper border of the rostral, which is often as deep as 
broad or a Httle deeper than broad (text-fig. 28 b) ; rostral " horn" 
with 3 to 5 transverse series of scales between the rostral shield 
and the apex. Supraciliary edge usually more prominent than in 
the typical form, sometimes slightly angular, foi'eshadowing the 
condition in V. raddii. Ventral shields 133 to 147. A more or 
less distinct dark blotch on the lower lip, involving 5 or 6 labial 
shields without interruption. Lower surface of end of tail yellow. 
Grows to 60 centimeti-es. 

Text-fig. 27. 

Side views of heads of a, Vipera ammoch/tcs, f. ti/pica (Feldkirchen, 
Garinthia), and b, var. meridionalis (Athens). 

Text-fig. 28. 


Front view of end of snout, showing the lepidosis. 
a. Fehikirchcn, Garinthia ; h. Athens ; c. Cocosu, Roumania. 

I submit these definitions to the consideration of herpetologists 
who may have the privilege of examining large series of speci- 
mens from any single district. The material at my command is 
still deficient in examples from Bulgaria, Turkey, and the Caucasus, 
and I may mention that the only two specimens from Roumania 
which I have been able to examine differ from both forms as here 
chai'acterized, and are on the whole intermediate between them. 
The rostral shield is deeper than broad, and the naso-rostral does 
not reach the canthus rostralis (text-fig. 28 c) ; one has two series 
of scales on the rostral " horn," the other has three ; supraciliary 
edge not very prominent ; ventral shields 155 and 153 ; no light 
bars on the lower lip ; lower surface of tail yellow. 


G. Notes on the Habits of the Hoolock. 
By Geo. Candler, M.B.Cantab. ^ 

[Received January 20, 1903.] 

The Hoolock {Ilijlolates hoolock) is one of the most interesting 
of the family Simiidte, and is perhaps not so familiar to natni'alists 
at home as are some other members of the family, as, owing to its 
extreme delicacy and the great ditiiculty experienced in keeping it 
alive in confinement, it does not often find its way into European 
collections. Even in the Calcutta Zoological Gardens it is difficult 
to keep Iloolocks alive for any length of time. They often 
succumb to pneumonia, or if they escape actual disease they mope 
and die from the eftects of confinement, or possibly from depri- 
vation of some article of diet Avhich in the wild state they have 
been accustomed to. I have ventured therefore to submit to the 
Society these short notes, made fi-om the point of view of a field- 
naturalist luther than fi-om a scientific aspect. 

The Hoolock is clothed all over with a fine soft hair, which, 
perfectly black in the male, in the female shows a greyish tint, 
especially over the back. This uniform dark colour is only 
relieved by narrow horizontal streaks of white hair above the eyes. 
The face, palms of the hands, and soles of the feet are devoi(l of 
hair, and here the black skin is smooth and finely wrinkled and as 
soft as the finest kid. 

The hallux and pollex have a flattened nail, the remaining digits 
have the nail laterally compressed and resembling a claw. 

Thei'e is no tail. Ischial tuberosities and cheek-pouches are 

Wlien the Ape is sitting, the vertebral column presents a single 
marked curve with the convexity backwai-ds. On the ground the 
Hoolock has a very characteristic gait. He goes along in a sort of 
sliambling waddle, with legs bowed and knees bent, the soles of his 
feet applied flat to the ground with the hallux widely abducted, both 
arms being carried upwards and extremely alxlucted as if to balance 
himself. He cannot get up any speed, and invarialfly swings up 
into the first tree he comes to, where his movements are suddenly 
changed from extreme awkwardness to extiuoi'dinary grace and 

He swings along to the thinnest part of a liough, or to the 
slender end of a bamboo, until it bends to his weight, then with a 
swing and a soi't of a kick-off he flies through the air, seizing 
another bough and swinging along it with the unerring accuracy of 
a finished trapeze performer. I fancy he does very little walking 
in the wild state, for I have never seen a wild Hoolock on the 
ground. Moreover, they are only found in the dense jungle where 
the ground is everywhere covered by tangled vegetation. It is 
puzzling to me why these anthropoids, being so entirely arboreal 

^ Coiiinmiiicatc'J liy F. (!. Paiisons, F.Z.S. 

188 MR. G. CANDLER ON THE [Feb. 17, 

in habit, should be lacking in such a useful appendage as a tail, 
I think, at any rate, that it points to the fact that the apes have 
been developed along a line distinct from the monkeys, the earlier 
traces of which line are yet to be discovered. 

The Hoolocks are extremely shy, and it is most difficult to 
watch them, as they are concealed by leaves high up in the tops of 
the bamboo- clumps or foi'est trees. You may hear their cries all 
round you as you I'ide quickly along a jungle-tract, but the 
moment you leave the path or look up at them there is a dead 
silence and scarcely a leaf stirs, until, tii'ed of waiting, you move on 

The cry of the Hoolock is a characteristic sound in the Cachar 
jungle. It is a very pleasing note, rising and falling in intensity, 
and reminding one somewhat in its rhythm of a pack of beagles 
giving tongue on a scent which is waxing and waning in strength, 
as a larger or smaller number of the band join in the choiais. 
It is heard chiefly in the early morning, then all through the heat 
of the day there is silence, but towards evening, as the sun sinks, 
you may hear it again. Hooloo ! Hooloo ! Hooloo ! with the accent 
on the Hoo syllable, is supposed to describe the sound, but it is 
really quite indescribable in writing. 

As in other species of apes, there is a special modification of 
the larynx, which acts as a sort of i-esonating-box, and helps (I 
suppose) to make the sound carry, as it does, long distances. There 
is also a peculiar arrangement of the upper aperture of the larynx, 
with its small and inadequate looking epiglottis, which more 
resembles the arrangement in birds than the leaf-like epiglottis in 

As, day after day, I have I'idden through the jungle, it has seemed 
to me that the Hoolocks work their ground systematically in 
their search for food, just as the planter plucks one section of 
his tea to-day and another section on a distant part of the garden 
to-morrow. Foi' I have found them filling the air with their cries 
along a particular sti'etch of jungle-road one day, whilst the next 
day not one was to be heard ; then, perhaps, a week later they ai'e 
back again in the same place. Living as they do in communities, 
they ai'e constantly on the move, and from what we know of their 
great intelligence, it seems to me highly probable that their move- 
ments are guided by very definite plans, and that very probably 
they have some sort of government system. 

There is a point about the Hoolock that strikes me as very 
extraoi'dinary, and that is the fact that he cannot swim. I had been 
told this by both natives and Europeans, but I confess I was 
somewhat sceptical about it until I tried experiments myself. We 
put a full-grown Hoolock into a big tank in 10 feet of water. He 
struggled helplessly, as a boy would before he learns to swim. 
He sank twice, with head thrown back and arms waving franti- 
cally, and we were obliged to rescue him almost asphyxiated and 
choking in the most human way. 


This weakness he shaves witli man, but I do not know whether 
(or not) it has been noted in the otlier antliropoids. 

It is a significant fact that the range of the Hoolock is l)ounded 
by two vast rivers, tlie Bralmiaputiu on tlie north and the Irawaddi 
on the soutli. It may well he that, with his natural aversion to 
water, these rivers have confined hiin to the comparatively limited 
stretch of countiy lie occupies. Tiuvelling high up in the jungle, 
he could swing easily across the ordinaiy sti'eanis which would 
come in his path ^\•ithout having to take to tlie water. The 
monkeys of India take readily to water, and it is a pretty sight to 
see them spring out from a lofty overhanging bough and drop, one 
after another, with a splash into the stream, and stiike out Ijoldly 
for the further bank. 

In Cachar, where these notes were written, the tea-plantei'S 
often keep Hoolocks for years, allowing them to run loose about 
the compound, and they are certainly the cleanest and most 
interesting pets imaginable, ofiering a \evy marked conti'ast in 
this respect to the red monkeys, which, chained to a pole, are so 
common a feature in Indian compounds. 

A Hoolock, to be tamed in this way, must be caught quite young, 
and not tied or shut up in any way. A native l)oy is generally told 
ofl:' to watch him for a few days, and to prevent him from bolting, 
but he soon learns to come down from the ti'ees for a plantain, 
and he will in most cases settle down to a solitary life, I'emaining 
about the .same compound for years. But chain him or restrain 
his libei'ty in any way, and he inevitably begins to mope and pine, 
and invai'iably dies in a, few weeks. It is strange that the calls of 
the wild Hoolocks, which he must hear almost daily all round him, 
do not tempt him to revert to his natui-al life as a member of a 
wandering community. I imagine a Hoolock, who attempted to join 
a strange band, would meet with a rough reception, anyway they 
never try to return to the jungle after they become tame. 

Several such tame Hoolocks I have had the opjioitunity of 
observing for some months past. Often they will be away up in 
the tree-to}is for days together, when nothing will tempt tliem 
down, ]:)ut when one chooses to be sociable he will come and sit 
on the arm of your chair at breakfast, and never reach or snatch 
things oft" the table: in fact his manners are unexceptionable, and 
he keeps his skin beautifully clean without that exaggerated 
parade of flea-hunting which makes the monkey tribe so oi>jection- 
able as pets. At sunset a-ou may see him settle down to sleep, 
jammed tight in a fork in a squatting position. In this semi- 
domesticated state I notice that the Hoolock seldom uses his 
voice. I suppose, leading a solitary bachelor life, he finds no 
necessity for chattering or calling. "With regard to the diet of 
the Hoolock, Dr. Blanford, the Indian natui'alist, gives a long 
list, including fruit, leaves, young shoots, spiders, insects, birds' 
eggs and young birds. But, it seems to me, the diet of such shy 
creatures must be largely a matter of conjecture, for no certain 


conclusions can be drawn from the habits of captured specimens, 
nor can we recognize as a rule substances in the stomach of shot 
specimens, as we can in the crop in the case of birds. My own 
observations lead me to believe that fruits and the succulent shoots 
of young bamboos and other trees form the bulk of their diet. 
They will certainly catch and eat certain spiders ; but I have 
invariably found them to refuse such insects as moths or butter- 
flies, perhaps because many such insects have a bitter taste. Eggs, 
too, I found they would not eat. If you give an insect or a small 
bird to a Hoolock he will certainly pull it to pieces, and possibly 
taste or bite it, but it by no means follows that it is one of the 
regular dishes he enjoys in his wild life. 

The following list of leaves and shoots which are eaten by the 
Hoolock is given by Anderson: — Morugo pterygosperitna, Sj^ondias 
manifera, jFicus religiosa, Beta vulgaris, Ipomcea reptans, Canna 

I hope later on to supplement these short notes with some 
anatomical observations on weight of brain relative to body, and on 
the number and depth of the convolutions. But this is a matter 
of time, for specimens are not very readily obtained. The Hindoo 
coolies, who form the bulk of the population in the tea-districts 
of Cachar, will never kill a Hoolock. The Kuki tribes in the 
Oachar Hills, on the other hand, kill and eat them, and regard 
them as somewhat of a delicacy, I believe. But even a Kuki finds 
it diificult to get a shot at these creatures, so shy are they and so 
active in their movements. 


March .'{, 1903. 

G. A. BouLENGER, Esq., F.R.S., Vice-President, 
in the (Jhair. 

The Secretary read the following report on the additions made 
to the Society's Menagerie in Fel)ruary 1903 : — 

The registered additions to the Society's Menagerie during the 
month of February were 57 in ninnber. Of these 8 were acquired 
by piesentation, 48 were received on deposit and 1 on approval. 
Tlie total numbei' of departures dui'ing the same period, by death 
and removals, was 105. 

Amongst the additions special attention may be directed to : — 

1. An example of Ouvier's Clazelle {Gazella citvieri), deposited 
by the Hon. Walter Rothschild, M.P., F.Z.S., on Feb. 9th. No 
example of this i-are North- Afiican Gazelle has been exhibited in 
the Gaidens since 18G7. 

2. A Tamandua Anteater [Taviandua tetradactyla) from South 
Ameiioa, received on approval on Feb. 12th. This is a very 
health}' and lively specimen of an animal which we have not had 
living in the Gardens for some yeai'S. 

3. A young nrde Chimpanzee {Anthropopithecus troylodi/tea) 
about a year old, deposited by Mr. J. C Lamprey, of the West 
African Regiment, on Feb. 18th. This animal is said to have 
been obtained at Kionko in the French Soudan, and was brought 
home from Sieriu Leone by the depositor. It is in very good 

4. A Fiilletl Lizai'd (C/damydosaui'us kingi) from Australia is 
the second example of this species received at the Gardens, the 
first having been pi-esented by Mr. Saville Kent some seven years 
ago. The piesent specimen was presented by IMi-. H. W. Fawdon 
on February 18th. 

The Secretary read the following extracts from a letter addressed 
to Mr. P. L. Sclater by Major C. Delme Radclifi'e from Uganda, 
and exhibited the skins of a Monkey {Cercocelms aterrhims) and 
an Otter {Lutra capensls) obtained by his collector J\Ir. Doggett, 
and sent home by parcel post, at the same time : — 

" I am sending you the skins of two monkeys — one completely 
prepared with bones, and the skin of the other, also skulls of both. 
The animals were both female. Doggett got them for me a few 
days ago, as I had sent him collecting by road here from Entebbe. 
This monkey is quite new to me. I shall be glad to hear from you 
if it is a new species. It is very intei-esting, and the skull has .some 
interesting features, for instance the small canines. The hair is 
curiously like a Chimpanzee's. Doggett tells me he thought from 
the noise they made that they were Chimpanzees at first. Is it a 
true Cercopithecus '? It interests me very much, and I shall be 

Proo. Zool. Soc— 1903, Vol. I. No. XIII. 13 

192 ME. w. E. DE wiNTON OX [Mar, 3, 

glad of your opinion. In the same parcel I enclose a Lake Otter 

Mr. F. E. Beddard, F.K.S., exhibited the mounted skin of the 
Greater Bird of Paradise [Paradisea apoda) that had lately died 
in the Society' Gardens. 

Mr, J. L. Bonhote, F.Z.S., exhibited a photograph of two tame 
Elephants in Ceylon accompanied by a baby one, which latter 
showed a considerable amount of hair, especially on the forehead. 
One of the old Elephants in the photograph, presumably the 
mother, also showed a ceitain amount of hair. 

With reference to a recent paper by Col. C. E. Stewart, dealing 
with the real home of the Tiger, Mr. J. L. Bonhote said that he 
had asked a friend of his, a Sanskrit scholar at Cambridge, as to 
whether there was a word foi' the tiger in Sanskrit ; the reply was 
in the aliirmative, the word being vydghra (or in the Pali form, 
vyaggho). This note was not brought forward against the fact 
of the original home of the tiger being in the north, but to support 
Mr. Thomas's contention that it had not spread south so recently 
as Col. Stewart was inclined to believe. 

Professor F. Jeffrey Bell, F.Z.S., exhibited a specimen of a 
Holothurian of the genus Actinopyga from shallow water off 
Zanzibar, in which there was not only an oral extremity with 
tentacles, as in the Ctocumaria planci described some ten years 
since by Prof. Ludwig, but also an anal extremity ; these additions 
(whether the result of gemmation or of fission cannot at present 
be said) do not occur in the same radius of the Holothurian. 

The following papers were read : — 

1. On a new Species of Pigmj^ Antelope of the Genus 
Neotragus from the Cameroons District^ W. Africa. 
By W. E. DE WiNTON, F.Z.S. 

[Received March 3, 1903.] 

(Plate XIX. & Text-fig. 29.) 

In a collection lately received from Mr. G. L. Bates from the 
Cameroons are two specimens, male and female, of a new form of 
Pigmy Antelope, adding a second species to the genus Neotragus. 
I propose to name this very interesting new form in honour of the 
collector, who has added so much to our knowledge of the fauna 
of the Camei'oons country. 








Neotragus batesi, .sp. n. (Plate XIX.) 

Size nearly half as large again as the Royal Antelope, jV. pyy- 
mceus. The colour ilarker and slightly more smoky, the feet not 
.so rufou.s, and the wliitish markings of the uudei-pai-ts not so 

The front of the face, crown of the head, ami neck are dark 
smoky- hi-own, the bnck and upperside of tail more rufous or bay- 
coloured. All the fur annulated with dark brown and bright bay. 

The t^vpe of this new species, a male, bears the following notes 
on the label attached by the collector: — " Efulen, Bulu Country, 
Kamai'un, 1500 ft. above sea. Caught in trap on edge of garden, 
15 Sept. 1902." Measureuieiits : head and Ijody 540 mm., tail 65, 
hind foot (without hoof) IGO, ear (inside from notch) 50. 

Dimensions of skull of male (text-fig. 29): — Greatest length 
107 mm.; basal length 95; greatest breadth 51 (below orbits) ; 
bi'eadth of brain-case 37'5 ; orbit to points of premaxilla? 48 ; 
length of nasals (middle line) 33 ; length of frontals 33 ; length 
of parietals 30; length of interparietals 13; greatest breadth of 

Text-fio-. 29. 

Skull of Neotrac/ us hate 

nasals 20 ; breadth in narrowest pai't 9 ; lengtli of horn (biolien) 
27, the peifect horn could not be less than 35 mm. in length ; 
distance between bases of horns 25 ; distance between tips 
{circa) 35 ; length of orbit 23 ; height of orbit 22 ; length of 
upper tooth-row 32; breadth of palate between pms.' 17, between 
ms.' 20, between ms.^ 21 '5. 

The dimensions of the skull of the female are practically the 
same as those of the male. 

The discovery of this new species extends the I'ange of the genus 
Xeotragus very considerably, for the Royal Antelope (i\\ pyymceus) 
is found onlv on the Gold Coast from Liberia to Lagos. 


194 MR. E. R. SYKES ON [Mar. 3, 

An examination of the skull somewhat modifies the definition 
of the genus as laid down in ' The Book of Antelopes ' by Sclater 
and Thomas ; for in this new species are found extra maxillo- 
premaxillary vacuities 9 mm. long and 3 mm. broad, very similar 
to those found in the genus Nesotragus of East Africa. The 
horns again of the members of this genus cannot be said to be 
" less than the diameter of the eye " in leng-th ; for the horns 
of the type of this new species, in their broken state, are longer 
than the diameter of the orbital cavity, and would undoubtedly 
measure half as much again in their perfect state. These horns 
are not "perfectly smooth," but show slight ridges or rings in 
their basal poi-tion. The skull is otherwise fairly similar to that 
of the Royal Antelope. 

There can be no doubt that Bates' Pigmy Antelope somewhat 
bridges over the difierences between this genus and the East- 
African Nesotragus, and practically reduces the distinguishing- 
characters to those of the horns alone. 

The horns of Neotragus are very small, practically smooth, and 
laid back on the head in a plane with the forehead ; while those 
of Nesotragus may be half as long as the head or more, strongly 
and closely ridged and directed upwards. 

2. On the Land Operculate Mollusca collected during the 
" Skeat Expedition " to the Malay Peninsula in 1899- 
1900. By E. R. Sykes, F.Z.S. 

[Received February 2, 1903.] 

(Plate XX/) 

The species of Land-Mollusks collected by the "Skeat Expedi- 
tion," though not very numerous, are of considerable interest on 
account of the welcome addition made to a fauna which is as yet 
but little known. They include : — 

Leptopoma aspirans Benson. 

Leptopoina aspirans Benson, Ann. Nat. Hist. ser. 2, vol. xvii. 
p. 229. 

Hah. Biserat, Jaloi-. It has been recoided from Bukit 
Pondong and the Kinta Valley, having been originally described 
from Tenasserim. 

Lagochilus kobelti, sp. nov. (Plate XX. figs. 13-15.) 
Testa modice tiinbilicata, turhinata, solidula, lirulis numerosis 
cincta, periostraco brunneo leviter induta ; spnra conica ; 
anfr. 5j, convexi; aper tur a modice ohliqua, suhcircularis, peri- 
stomate dtiplici, incrassatulo et suhreflexo, juxta insertionem 
breviter sed distincte inciso. Alt. 6"8 ; diam. max. 7'0 millim. 
Hab. Biserat, Jalor. 

1 For explanation of the Plate, see p. 199. 


J.GT-eeTxael.etifK 18. 19. Mi:r.terix Bros imp. 



The species of Lagochilus recorded from the Malay region are 
very puzzling and I think Dr. Moellendorft" Wiis qixite right in 
describing as L. rollei the form that I noted (under the name of 
L. toivnsendi) from Kelantan. The nearest ally of L. kohelti 
appears to be L. toivnsendi Crosse ; I have not seen an authentic 
specimen of that species, but have compared the form now described 
with the description and figures given by Crosse and with a 
specimen collected by Herr Grubauer, from whose collections 
Dr. Moellendorft" has recorded L. tovmsendi as the only species 
foinid. The shell I now name is a trifle smaller and more 
elevated in proportion to the breadth, and the umbilicus is 
narrower. It may be noted that the reference to Crosse's original 
paper in the ' Journal de Conchyliologie ' should be to p. 200 and 
not p. 208 as given by De Morgan and Dr. Moellendorff in their 
papers on the Pei'ak fauna. 

I have named the form after Dr. Kobelt as a trifling recognition 
of his recent study of the Cyclophoiida?. 

DiTROPis CAVERN^E, sp. nov. (Plate XX. figs. 17-19.) 

Testa depresso-conoidea, late tombilicata, olivacea, tenuis, glabra ; 
s]nra medioa'iter elevata, apice eroso, sutiiji'a impressa ; 
anfr. 4 (?), convexi, ulthmis antice vix descendens, carinatus, 
carinis chmhus su])ra pei'iphe7-iavi, unica ad, jjeriphet-iain, et 
plurimis in regione umbilicali ; apertura subovalis, peristotnate 
incrassatido. Alt. 2*2 ; diam. max. 1*7 milliiti. 

Hab. In a cave, Biserat, Jalor. 

A single specimen only. 

Cyclophorus malayanus (Benson). 

Cyclostoma malai/anitm Benson, Ann. Nat. Hist. ser. 2, vol. x, 
p. 269. 

JIab. Gunong Inas, Perak. 

Recently when cataloguing (J. Malac. ix. p. 61) a collection of 
shells from Kelantan, I gave the names of Cyclojyhorus saturnus Pfr. 
and borneensis Mete. : both the forms then recorded have occurred 
in the present collection, and I have therefore again considered 
the identifications. Both, according to my present view, are 
erroneous, and the group is a very difficult one. The present 
species, which I regard as a form of C. malayanits, was then named 
G. saturnus, and the next species was called C. borneensis. 

Cyclophorus tuba (Sby.). 

Cyclostoma f^66rt Sowerby, Pi'oc. Zool. 8oc. 1842, p. 83. 
Hab. Gunong Inas, Perak. 

See remarks under the last species ; probably the C borneensis, 
recorded by De Morgan from Perak, also belongs to this species. 

Pterocyclos subalatus, sp. nov. (Plate XX. figs. 1, 2.) 
Testa late umbilicata, orbiculato-dejiressa, lineis incrementi notata, 

196 ME. E. R. SYKES OK [Mar. 3, 

hrunnea, strigis cornels picta, fascia unica nigro-hrunnea ad 
peripherimn ornata ; anfr. 4|, mediocriter crescentes, coiivexi, 
sut'iora bene notata separati ; apertura siobcircularis, peri- 
stomate indistincie duplicato, ala parva antice aiigidato. 
Alt. 8 ; diarn. max. 1 6 millim. 
Hah. Gunong Inas, at 5000 feet, 

I thought at fii\st that this might be a form of P. hlandA Bens., 
but it appeal's to be smallei', more elevate and not so widely 
umbilicated, and to difler in colour. In the two specimens that I 
have seen the lip is diiplicated only on its outer mai-gin, and the 
wing is small and thin. 

Rhiostoma jalorensis, sp. nov. (Plate XX. iigs. 6-8.) 

Nearly related to R. housei but larger, slightly more depressed, 
and with the whorls not so tightly coiled. The tube is long and 
large, reaching the body of the shell, and is bent slightly back- 
wards at the junction. The disjoined poition of the last whorl is 
much longei' than is the case in E. housei, as will be seen from 
the figure, and is more descending. The colour resembles that 
of B. housei, and a dark band is usually present at the periphery. 
Operculum as usual. Diam. max. 29 millim. 

JIab. Limestone Hills and Oaves, Biseiat, Jalor. 

I have been in some doubt as to whether this is not a local race 
of 7?. housei, but the difierences are constant in the specimens 
examined and, I think, are of specific value. The porcellaneous 
appearance of the operculum, so often seen in this group, appears 
to be due to the wearing down of the outei- layers. 

Rhiostoma, sp. 

Hah. Kwala Aring, Kelantan. 

A single specimen which agrees well with R. jousseaumei 
De Morgan, save that the tube is placed a little further back from 
the mouth, and is thinner and moi'e cylindrical. It appears safer, 
however, to await fui'ther material ere describing this form. 

Opisthoporus penangensis Stol. 

Opisthoporus penangensis Stoliczka, J. As. Soc. Bengal, vol. xlii. 
pt. 2, p. 265, pi. X. fig. 7. 

Hah. Kwala Aring, Kelantan. 

A single specimen, which I refer to this species with some doubt. 

Rhaphaulus ascenders, sp. nov. (Plate XX. figs. 11, 12.) 

Testa pupoidea, angxiste ^mnhilicata, fusco-purp)urea, dense costu- 
lato-striata, sp>ira bene elevata, apice obtuszdo ; anfr. 6, plano- 
convexi, p)emdti')nus -gibbosus ; apertura suhcircidaris, intus 
pallidefusca; peristoma jxdlide corneum, exjjansitm et re/lexum, 
marginihus callo junctis ; tuhulus brevis, incrassatus, a sutidra 
oblique ascendens. Alt. 18'5; diam. max, 9*5 millim. 

Hah. Patalung. , 


A single specimen "fi-om rotten wood." Recalling in shape 
R. jjerakeiisis )Smith, the pi'esent species is larger and stouter, and 
the tube slants obliquely upwards instead of descending ; as com- 
pared with R. lorraini Pfr., which also has an ascending tube, the 
form is not so cylindrical, the whorls are flatter, and the size is 
much greatei'. 

Rhaphaulus perakensis Smith, var. jalorensis, n. var. 
(Plate XX. figs. 9, 10.) 

Shell very similai' to R. perakensis Smith, but a little thinner 
and palei- in colour, and the tube is bent along the outer lip, being 
attached to the lip, a.nd not diverted behind it as in typical 
R. perakensis. The tube is broad and short. This form may pi'ove 
to be a distinct species, but bearing in mind the great vai'iation 
shown by Col. Godwin-Austen (Moll. India, vol. i. p. 202, pi. xlvii.) 
to exist in R. blavfordi Benson, and also considering how little we 
know of the group, most forms having been described from very few 
specimens, I have thought it wiser to give only a, vai'ietal name. 

Hab. Bukit Bisar, on the borders of Jalor, altitude 2000 feet. 

A single specimen. 

Schistosoma anostoma (Benson). 

Cyclostoma anostoma Benson, Ann. Nat. Hist. ser. 6, vol. x. 
p. 269. 

Uab. Belimbing, Ligeh. 


Cyclostoma sectilabriim Gould, Boston Journ. Nat. Hist, vol, iv. 
p. 459, pi. xxiv. fig. 10. 
Hab. Ulu Selama, Perak. 

PuPiNA Lowi De Morgan. 

Pupina loivi De Morgan, BuU. Soc. Zool. France, vol. x. 1885, 
p. 414, pi. vii. fig. 3 {loitri on plate). 
Hab. Gunong Inas, Perak. 


Pupina aureola StoHczka, J. As. Soc. Bengal, vol. xli. pt. 2, 
p. 267, pi. X. figs. 11, 12. 

Hab. Jalor, a single specimen. 

Agrees well with Stoliczka's figure 1 2, but his figure 1 1 looks as 
if it might belong to a different species. 

Alyc^us thieroti De Morgan. 

Alycceus thieroti De Morgan, Bull. Soc. Zool. France, vol. x. 
1885, p. 403, pi. viii. fig. 6; Moellendorfl; Proc. Zool. Soc. 1891, 
p. 342. 

Hab. Belimbing, Ligeh, a single specimen. 


Alyc^us diplochilus Moellendorff. 

AlyccBus diplochilus Moellendorff, J. As. Soc. Bengal, vol. Iv. 
pt. 2, 1886, p. 313 ; Proc. Zool. Soc. 1891, p. 342, pi. xxx. fig. 8. 
Hah. Cave near Biserat, Jalor. 

AlyCjEus conformis Fulton. 

Alycceus covformis Fulton, Ann. Nat. Hist. ser. 7, vol. ix. 1902, 
p. 68. 

Hab. Kwala Aring, Kelantan. 

Alyc^us perakensis Crosse. 

Alycceus perakeoisis Crosse, J. Conchyl. vol. xxvii. 1879, p. 206, 
pi. xii. fig. 7. 

Hab. Biserat, Jalor, amongst rocks. 

Opisthostoma annandalei, sp. nov. (Plate XX. figs. 4, 5.) 

Testa conico-pyramidalis, perforata, ])allide riifescens, spira bene 
elevata, apice obtuso ; anfr. 7, convexi, primi Iceves, reliqui 
distante lamellati, ultimios constrictus, retroversus, ascendens, 
conspicue solutus ; apertura rotundata, peristomate leviter 
incrassato. Alt. 2"2 ; diam. max. 2"8; diam. niin. 1*6 millim. 
Hab. Jalor. 

A single specimen, found by Mr. Annandale in debris on the 
floor of a cave. Related to 0. laidlawi Sykes, from Kelantan, but 
the pi'esent species is larger, and the spire is much more produced, 
thus becoming more cylindrical in appearance. 

DiPLOMMATINA SKEATI, sp. nov. (Plate XX. fig. 3.) 

Testa sinistrorsa, vix rimata, ovato-fusiformis, aurantio-rufa, 
solidiuscida ; anfr. 6^, convexi, p7-imi loives, reliqui costidis 
p)arvis subremotis regidariter scidpti ; sutura bene impressa ; 
apertura fere circtdaris, peristomate incrassatido, bene reflexo, 
subalato ; lamella columellaris paj-va. Alt. 3*1; diam. max. 
r9 millim. 
Hab. Gunong Inas, Perak, at about 5000 feet. 
Only a single specimen of the shell, the most salient features of 
which are the inflated whorls, deeply cut suture, and fine, regular 

DiPLOMMATINA LAIDLAWI, sp. nov. (Plate XX. fig. 16.) 

Testa sinistrorsa, rimata, ovato-fusiformis, cornea- albida, apice 
ritbello ; anfr. 5|, convexi, primi Iceves, reliqid costulis parvis 
remotis regidariter scidpti, interstitiis sub lente dense spiraliter 
striatis ; apertura quadrato-ch-cidaris, j^^^i^iomate duplici, 
expanso, margine coluinellari sinuato, subalato; lamella 
columellaris minima, indistincta. Alt. 2; diam. max. 
1 millim. 

Hab. Gunong Inas, 3000-4500 feet, Perak. 

Found " on the under surface of dead leaves, young with adults." 


Much smaller than the last species, the costula^ more distant, and 
densely marked with minute spirals. I can find no trace of these 
latter in T). skeati, hut the single specimen was not taken alive. 

Georissa moxtekosatiaxa Godwin-Austen k. Nevill. 

Georissa monterosaliaua Godwin- Austen & Nevill, P. Zool. Soc. 
1879, p. 729, pi. lix. %. 6. 
Hab. Pcrak. 


Figs. 1, 2. Fteroci/cJos suhalatus, p. 105. 
3. Diplommatina skcati, p. 198. 
4, 5. OpiaHioslnina annandalei, p. 198. 
6,7,8. R/iiosfimiti Jalorensis, p. 196. 

9, 10. Rhaphaidus perakfusis, var. Jalorensis, p. 197 
11, 12. Rhaphauliis asccndons, p. 196. 
13, 14, 15. Lngorhilm kohciti, p. 194. 

16. Diplommatina laidlawi, p. 198. 
17, 18, 19. Ditropis caver nee, p. 195. 

3. The Significance of the Callosities on the Limbs of the 
E<pdd(v. By R. Lydekker, F.Z.S. 

[Received Februaiy 5, 1903.] 

The question as to what structures in other mammals are 
represented by the callosities on the inner sides of all the limbs of 
the Horse, and those of the hind-lindis of the Kiang, Ass, and 
Zebras, is one which has attracted the attention of many naturalists, 
nearly all of whom appear to l)e in accord in i-egarding them as 
vestigial structures. The late Sir W. H. Flower, for instance, in 
his work ' The Horse ' expressed the opinion that these structures 
ai'e degenerate glands ; pointing out at the same time that the 
so-called ei'got on the hinder aspect of the horse's pastern appears 
to represent one of the pads, or cushions, which aie still functional 
in the foot of the Tapir. 

In his volume on " Mammalia " in the ' Cambiidge Natural 
History'', of which the preface is dated Feliruary 1902, Mr. 
Beddard gives a qualilied support to this gland-theory ; stating in 
one passage that the equine front callosities probably coirespond 
to the carpal glands of several other mammals, although on 
another page their glandular nature is questioned. In an 
apparently later communication " the same gentleman suggests 
that the callosities on the fore-lind)s of the EquAdjce may represent 
a carjial sense-organ, vestiges of which he believes to survive in 
the caipal bristles of the Dassies. The degeneration of such an 
organ would, it is urged, very likely result in the formation of 
sti'uctures resembling those under consideration. 

1 Pp. 12, 13, and 210. 

2 Proc. Zool. .Soc. 1902, i. p. 135. — I am indebted to Mi-. Heddard liimself for 
directing my attention to thi.s passage. 


There has, however, long existed an idea that the equine callo- 
sities are i-emnants of a vanished toe. Somewhat analogous to 
this idea is a theoiy, which has been advanced in a papei* lately 
read befora the Royal Society of Edinburgh by Prof. Ewart \ 
That genl:leman, it appears, has based his investigations to a 
very great extent on the condition obtaining in the foetus ; and 
has arrived at the conclusion that the callosities in question have 
nothing to do with glands, but that they i-epresent certain foot- 
pads of polydactyle mammals. According to Prof. Ewai't's view, 
the foi'e- callosity of the Hoi'se is homologous with the supplemental 
pad on the fore-foot of the Dog, while the hind callosity (which is 
wanting in all existing Equidce save the true Horse and the so- 
called Eqitus jjrsewalskii) corresponds to the hinder plantar pad 
of the Banded Anteater (^Myrmecohiusfascicdus). Such correlations 
will, I venture to think, scarcely be taken seriously by the great 
majoiity of zoologists ; and I shall thei'efore confine my attention' 
to the question whether the identification of these callosities with 
foot-pads geneiully is well founded. In all cases my observations 
are confined to the condition obtaining in adult animals. It may 
be added that I purposely brought these observations to the 
notice of the Society before the publication of Prof. E wart's paper, 
since I treat the subject from a different standpoint. 

In the first place, the callosities on both limbs of the Horse are 
situated on the inner surface, whereas, if they represented 
vestigial foot-pads, their position should be, i:)ri'mcii facie, on the 
hinder aspect, as is the case with the ergot. It might, indeed, 
be argued that they have changed their original position, but 
of such a shifting there is no evidence in the adult. A second, 
and perhaps more impoi'tant, objection to the foot-pad theoiy may 
be drawn from the fact that the callosities in the fore-limb are 
situated above the so-called knee-joint (carpus), and are therefore 
altogether higher up than any of the foot-pads of plantigrade 
mammals. Tiniest;, therefore, another shift of position has taken 
place, the fore-callosities do not represent foot-pads. This argu- 
ment, it may be mentioned, was used by Sir W. H. Flower to 
disprove the theory that the callosities aie remnants of the lateral 

The hind-callosities, on the contrary, are situated a short distance 
below the joint of the hock (tarsus), and ai-e therefore on a part 
of the limb, albeit on its inner side, which is included in the foot 
of a plantigrade mammal. If, however, the front callosity be 
regarded as serially homologous in a general sense with the hind 
one — and this is an integral part of Prof. Ewart's theory, — it will 
be evident that in the event of the former not being a foot-pad, 
the same will hold good for the latter. 

A third, and perhaps stronger, objection may be urged against 
the foot-pad theory. On the assumption that the callosities of the 
existing Eqiddce are vestiges of foot-pads, it is clear that these 

1 See ' Nature,' vol. Ixvii, p. 239 (1903). 


btructureb luuht have existed in the ancestors of that family ever 
since the time when these ancestoi-s were plantigrade. But, so far 
as I know, no ungulate was ever wholly plantigrade in lioth feet; the 
nearest npproacli to this condition nlitaiuing in the Lower Eocene 
Corj/phodoii, in which the hind-liml) was wholly plantigrade, while 
the front one was partially digitigrade. It has thus to hea.ssuined, 
on the foot-pad hvpothesis, that the front callosities of the Horse 
have been fuuctioidess structures from a period antedating the 
evolution of the Ungulata. 8uch a persistence, on exposed parts 
of the body, of a wholly fuuctioidess structui-e seems very im- 
probal)le, especially when the modifications are borne in mind 
which, on this hypothesis, the horse-line must liave undergone 
since the time when the callosities were functional structures. 
Perhaps the case of the ergot may be cited against this argument ; 
but it should be remembered that this structuie certainly acted 
as a functional pad at a much later stage of evolution than could 
possibly have been the case with the callosities. 

Having now stated what appear strong objections, so far a.^; the 
adult is concerned, against correlating the callosities of the Horse 
with the foot-pads of polydactyle mannaals, it remains to consider 
whether they can lie identified with any othei' structures. Those 
familiar with the morphology of the Cervidce, will be awai'e that a 
certain niunber of representatives of that family — notably the lein- 
deer, the White-tailed Deer, the Mule- Deer, and, in a rudimentary 
condition, the Elk, — are furnished on the inner side of the hock 
with a glandular tuft corresponding very closely in situation with 
the hind-callosity of the Horse. In fact, the only difference in the 
position of the two structures is that the tarsal tuft of the Deer 
in question is placed rather lower on the hock. From the fact of 
its occurrence in Deer so widely separated fiom one another as 
are the species mentioned, it seems evident that the tarsal gland 
(which is doubtless a scent-organ) is a very ancient structure, 
which was present in all the ancestors of the gi'oup, but has been 
lost, probably from disease, in the great majority of Old World 

Judging from their position, there would seem to be a certain 
probability that the hind- callosities of the Horse and the tarsal 
gland of the Deer are homologous structures. 

With regard to the homology of the fore-callosity of the Eqv,id(T, 
it may be mentioned that man}' Gazelles have tufts of hair 
(" knee-brush&s ") at the knee (carpus), which are probably 
glandular in origin. And it is possible (if the suggestion with 
regard to the hind-callosities hold good) that these may represent 
the fore-callosities of the Horse, for there seems no good reason 
why the position of a gland should not have somewhat shifted in 
two widely separated groups of mammals. Then, again, we have 
the carpal bristles of cei'tain mammals, such as the Ooatis and 
Dassies, already refei'red to as lieing regardeil by Mr. Beddard as 
the remnants of a " scent-organ," — a. structure probably not far 
removed in its nature from a gland. The occurrence of these 


bristles in the Dassies (Procavia) is very important. Mr. Beddard 
states that these are the only ungulates in which he has found 
these bristles. Carpal callosities ai-e, however, described by 
Dr. W. Leche^ as occurring in Wart-Hogs [Fhacochoerits); although 
they are stated by their describer to be acquired, and not primitive 
structvires. Whether the latter statement is calculated to modify 
Mr. Beddard's opinion with regard to the nature of the carpal 
bristles in the Dassies, I am, of course, unable to say. Of special 
impoi'tance is the occui'rence of bristles in these structures, since, 
even if hairs be found to exist on the callosities of foetal Eqiddoe, 
this would be no bar to the svipposition of their glandular nature. 

As regards the structure of the callosities themselves, it may be 
noted that in the Horse both pads are of a distinctly warty nature, 
and that the hind pair are certainly in a more decadent condition 
than the other, being in fact on the verge of disappearing. In 
the Zebras, on the other hand (in which the hind one has been 
lost), the fore- callosity is larger and much less warty and also 
situated higher up. In dried skins it is, in fact, much more like 
the pale glandular patch of skin below the ear of a Reedbuck ^. 
In this connection we have to bear in mind not only Mr. Beddard's 
observations alluded to above, but likewise others by Mr. Bland 
Sutton'^, in which it is pointed out that in certain Lemurs decadent 
glands are actually converted into bunches of spines, which are 
practically almost the same as warts ; that is to say, they are 
hypertrophied growths of somewhat abnormal dermal tissue. 
Hence there seems no primd facie reason why the callosities of 
the Equidce should not be decadent glandular structures, the 
decadence being more marked in the two pairs of callosities of the 
Horse than in the single pair of the Asses and Zebras. 

There is, however, another point which may have an important 
bearing on the subject. Fiom the presence of a depression in 
the skulls of Hipparion, Hippidium, &c., it is evident that 
primitive Horses were furnished with face-glands comparable to 
those of Deer ; such glands probably having a function somewhat 
analogous to that of the scent -glands on the limbs of the latter. 
If, then, the existing Equidce have got rid of their face-glands, as 
being (perhaps on account of change of habit) useless, it is con- 
ceivable that, for the same reason, they may have also discarded 
their limb-glands. 

If these suppositions (and they are but suppositions) be well 
founded, it follows that a tarsal and a carpal gland must have 
existed in the common ancestors of the Horses and Deer ; that is 
to say, in the common stock of all modern Ungulates save the 
Elephants and perhaps the Dassies. And it may be ui'ged that if 
this were the case, traces of such glands ought to be met with in 
Tapirs, Rhinoceroses, Pigs, Hippopotamus, &c. So far as I am 

1 Biol. Centralblatt, vol. xxii. p. 79 (1902). 

2 It would be important to examine the histological structure of ;the callosity in a 

3 Proc. Zool. Sec. 1887, p. 369, 


aware, tlie only instances of structures wliicli can be regarded as 
at all approaching this nature are tlie carpnl Ijristles of the 
Dassies and the carpal callosities of Wa.rt-Hogs, the latter of whicli, 
as already mentioned, ai-e considered to be of niodei-n origin. No 
tiace of any structure compaiable with the hind-callosity of the 
Hoise has, so far as I am awai-e, ever been detected on the tarsus 
of any of the above-mentioned Ungulates. 

If an objection of this nature be legai'ded as fatal to the gland- 
theoiy (or sense-oi-gan- theory, foi' I i-egai'd the two as practically 
identical) of the origin of the equine callosities, it will, I thiidc, 
be still more so to the foot-pad hypothesis, since short-limbed and 
polydactyle mammals ought to have I'etained traces of ancestral 
foot-pads for a gi'eatei' period tlian long-limbed moiiodactyle forms 
like the Kqukhe. 

In conchision, I may state that it has been my object, not so 
much to attempt to show what the equine callosities repi'esent, as 
to demonstrate, from pahi?outological considerations, the impro- 
bability of their being vestigial foot-jjads. 

P.S. — I am informed that if a callosit}' be pared down, and a 
finger moistened with the I'esnlting exudation held to a horse's 
nose, the animal will follow nnywhei'e. If tins be true, it afibrds 
strong testimony in favour of the gland-theory. 

4. Note on some Remains o£ Strut /i to karat /leodoris Maj. of 
the Island of Samos. By Rudolf Martin, of Basel 

[Received Pebruaiy 11, 1903]. 

(Text-figures 30-34.) 

In the Catalogue of the collection of Mr. Barbey at Valleyi'es 
s./Orbe (Switzerland), published in 1894 l)y L)r. Forsyth Major'-, 
besides a great number of mammals there is mentioned the femur 
of a ratite bird, which, no doubt, belonged to the genus Slruthio 
(Dr. Major could scarcely find any diflerence). Dr. Major, 
considering the geological age of the deposits in whicli the bone 
had been found (Upper Miocene), and recognising that it belonged 
to a form dillerent fi'om the I'ecent Ostrich, created the navf species 
Strathio karatheodoris. 

Some time ago, Dr. Major received the fragment of a pelvis 
from the Museum of the Vienna University (found in the same 
place as the femur), which probably belonged to the same species, 
and which is the subject of the following remai'ks. 

I have to thank Dr. Major for having entrusted me with the 
study of this pelvis and for having placed two photographs of the 

' Coiniiuuiicutcd by Dr. C. I. Forsyth Major, F.Z.S. 

- Le (u-^cineiit (issifi'iv do Mitylini et Cataloarue d'ossements fossiles ricueillis a 
Mityliui etc., Lausanue, 1894. 



pMar. 3, 

femur (rostral and cavidal aspect) at my disposal, on which the 
following description is based. 

Evidently this femur (text-figs. 30, 31) belonged to a large 
struthioiTS bird, as it differs little from the same bone of a modern 
Ostrich. The size of the femur is somewhat larger than that of 
the same bone of recent Ostriches at my disposal, but very probably 
there Avould be no difli'erence between a, large specimen of Struthio 
camehts and the fossil. 

Text-fig. 30. 

Text-fiff. 31. 

Text-fig. 30. — Right femur of Struthio karatheodoris. Caudal view. 5: nat. size. 
Text-fig. 31. — „ „ „ „ Rostial view. \ nat. size. 

The following table gives the results of a compaiison between 
the fossil in question and that of a medium-sized modern 
ostiich : — 

Length of Length of caput Smallest Width of 



-(- coUum. 



Str. karatheodoris 




12"4 mm, 

Str. camehis 




11-5 mm, 

The measurements of the caput -f collum have been taken 




from the apex of the caput to tlie margin of the trochanter, and 
that of the troclilea in the direction of its greatest expansion.' 

A close stud}' of the bone, liowever, revealed some features in 
which the fossil differs from the recent Ostrich, on the sup- 
position that the photograph is not distorted. 

In the fossil, the neck supporting the head of the femur is much 
stouter than in the recent species and is much less constricted. 
This is best seen in the form of the distal border of tlie collum. 

^ Although nieasurements may include an error of, say, -L, they 
give an idea of that feature, because the difference between the 
two species exceeds that error. The only measurable line is the 
plummet in the apex of the hea.l to tlie liiiext uspera, divitlin^'- 
the rostral surface of the neck from that of the bone itself. ° 

Width of the neck = 1 . 
StriitJi'io cameliis = A. 

„ karatheodoris = ^ 

It seems to me that the groove in wiuCli the Hguuieiituin teres 
is inserted is shallower in the fossil than in ,Struthlo cumelas. It 
may be that the photograph gives a wrong impression, and I think 
it would be better not to regard this feature as a specific one. 

Text-fiff. 32. 

"I' S/rnfh;,, k:,r.ith,'odoris. \ vmU'aX view 
« & 6 = true -acral vertehi-.u. i.. ii., & iii. = first tin 
,', iiat. size. 

ee post sacral vertebra?. 




The trochanter and the li7ie(e asperce, of the proximal end of the 
femur are not to be distinguished from the same parts in the 
recent Osti'ich ; even the pneumatic foiumen in the caudal surface 
is found in the same place. 

The middle pai-t of the bone is just as in S. cainelibs, nd the 
distal end diffei-s only in one particular : the fossa intercondy- 
loidea is much elongated, so as to form a slight valley in the distal ^- 
oi" I of the I'ostral suiface of the femur. In the caudal surface, 
immediately above the proximal border of the trochlea, there is 
also a funnel-shaped pneumatic foi'amen which enters the bone 
obliquely (fi'om the mesial border of the bone, and is directed 
towards the middle of the axis of the trochlea). As in the recent 
Ostrich, there is a tuberosity" between the wide orifice of that 
foramen and the mesial edge of the caudal surface. 

The pelvis (text-figs. 32-34) difl^ers in some respects rather 
considerably from that of the recent Ostrich ; but the individual 

Text-fig. 33. 

Pelvis of StrutMo karatheodoris. Dorsal view. 

variation met with in the recent foim, and very piobably also in 
the fossil, shows that the difliierence is much less than it seemed to 
be at first sight, and very probably the extreme variations of 
the two forms closely approach. 


The pelvis at my disjiosal is lepresented only by a fragment 
from the last pre.9rtc;-rtZ to the t\\ivi\ postsacral vertebra (fi vertel)ra', 
the first and the last incomplete), and in conneetion with it tlie 
corresponding 2)arts of the ilia, *.e.the ncetabularaiid immediately 
postacetabular i-egion. The acetahida themselves are not entii-ely 
preserved, but only the caudal liorchn' of that on the left is quite 
intact, and the antitrocltaiiter is broken away on both sides, so 
that on the left side only its outlines can be deteiinined. 

Text-fi^. 34. 

Lateral view of the pelvis of Strufhio karatJieodoris. \ nat. size. 

In the principal and most characteristic features, the fossil 
does not difi'ei- from *S'. camelus, and there can be no doubt that 
the ratite bii'd of Samos belonged to the same genus. 

In the following featui'es the fossil pelvis agrees with oi- differs 
from that of the recent Siruthio : — 

i. The pelvis is compressed from both sides in the same mannei" 

as in StrutJtio camelus (though tlie comi)ression is a little less 

marked), so that the dorsal as})ect of it is much as in the 

recent form. 
ii. The sacral vertebra', of the fossil ai'e much stouter. The 

same iri-egulai- sculptures occui', foi-ming lough longitudinal 

iii. The centra of the two " true sacral vertebra', " are, compai-ed 

with the following centia, nari-ower, but better rounded 

than in the recent Ostrich, 
iv. The acetabulum has quite the same position as in aS'. camelus. 
V. The plane between the venti'al border of the antitrochanter 

and the dorsal edge of the foramen obturatorium is— in 

comparison with >S. camelus — turned foi'ward, i.e. it slopes 

more rapidly towards the acetabulum. 
vi. The outlines of the antitrochanter ai'e just as in S. camelus, 

and its size is also the same. 
vii. The ilia of the fossil and the recent Ostiich are closely 

similar. There is no difteience in the dorsal view of the 

pelvis exceeding possible individual variation. The vertical 
Proc. Zool. Soc— 1903, Vol. I. No. XIV. 14 


distance between the antitrochanter and the lateral edge 
of the area dorsalis is relatively longer in the fossil than 
in most of the recent medium-sized Ostriches ; but on one 
side this distance cannot exactly be determined in the fossil 
(because the antitrochanter is broken away and the edge 
of the area dorsalis is mu^ch rounded), and on the other side 
the individual variation in this regard in S. camelus is very 
great. The largest individuals of this species differ but 
little from the fossil, and very probably the fossil pelvis at 
my disposal must be referred to an adult and rather large 

The distance between the dorso-mesial borders of the ilia is 
somewhat shorter in the fossil than in most of the pelves of 
aS'. camelus with which I have compared it. But it is in this respect 
that individual variation has the greatest amplitude, and in the 
largest pelvis of ^S'. camelus at my disposal this distance is equal 
to that of the fossil. 

/S. Icaratlieodoris. S. camelus. 

Costal process of the 1st '' true sacral 
vertebra" to the distal border of the 
2nd postsacral vertebra 7*8 cm. 9*2 cm. 

Width of the centra of the " true sacral 

vertebrae" 2-5 1-8 

Width of the centrum of the 2nd postsacral 

vertebra 2-8 2-3 

Greatest width of the dorsal surface of the 

pelvis 12 12-2 

Plummet in the dorsal border of the anti- 
trochanter to the median plane ca. 1 2 12 

Plummet in the dorsal border of the anti- 
trochanter to the plane of the dorsal 
surface of the pelvis ca.6'5 4 

The results show that there are great differences in the pro- 
portions of the pelvis of the two species which justify a 
separation of the fossil from the recent Ostrich. The individual 
variation, however, does not allow us to place the two forms in 
different genera, because it greatly reduces the differences above 

I have said that the separation of the two Ostriches in different 
species was justified. On the other hand, the question arises, 
whether or no the differences between the two forms be not the 
consequences of a pressure acting during the great tectonic trans- 
formations in the region between Asia Minor and Greece. A 
study of the other fossils of Samos, however, gives no evidence of 
such a force ; there is only a lateral compression to be seen, a result 
of the pressure of the weight of the more recent strata. 

These considerations add to the importance of the features which 
I have selected for special comment, Avhilst on the othei' hand other 


featm^es lose their systematic value. But I have sliown that the 
latter had really already lost their importance because of their 
individual variation. 

The species Struthio karaiheodoris Maj. is therefoi'c to he kept 

What are the relations between ^S'. karatheodoris and ^S'. asiaticus 
from the Siwalik Hills? I can now give further information 
on *S'. asiaticus. A comparison of the fossils with the figures on 
the unpublished plate R of Falconer's ' Fauna Anti(|ua Sivalunsis,' 
and those accompanying Davies' ^ paper and Lydekker's 'description, 
did not reveal any differences, except that the drawings in 
Falconer's Atlas are much better than the others. Davies found 
that the principal difference between S. asiaticus and >S'. camelus lies 
in the greater stoutness of the cervical vertebra) in the former ; 
and Lydekker kept the two species sepai\ate onlj^ on account of 
this feature. The answer to the question, whether the greater 
stoutness of the sacral vertebrae on one side and tliat of the cei'vical 
vertebraj on the other indicates a special i-elationship, cannot be 
given now. There is no evidence for such relations ; and I, for 
my part, eonsidei- this character in the fossil forms (compared with 
the modern Ostriches) only as more primitive. 

There may be i-eason to unite the two forms in one species ; but 
considering their different geological ages', I think it will be better 
not to do so. But there can be no doubt that Struthio asiaticus 
is in direct relation with >S'. karatheodoris, i. e. that the former is 
a descendant of the latter, as Dr. Major' supposes the whole 
Siwalik fauna to be a later and transformed generation of the upper 
Miocene fauna of Pikermi and Samos. 

^ It is noteworthy therefore that S. karatheodoris and S. asiaticus 
give us no evidence for a specialisation of the Struthionidte in 
Southern Eurasia", and a consequent emigration into Southern 
Europe, S}a-ia, and Africa, but support rather the view that the 
order of migration took place in the opposite direction. However, 
the genesis of the Struthionida; cannot, as Burckhardt ' is inclined' 
to suppose, be associated with the Miillerornithida- ; for the 
geological age of the former is opposed to such an hypothesis. 

By the discovery of a Struthio in the island of Samos, the eo-o- 
on which the species of Struthio chersonensis Brandt ' has been 
based is of some interest. Though the circumstances surrounding 
its disco\'ery may be somewhat obscui-e, yet the size and proportions 
show that it cannot be the egg of a modern Ostrich ; tliat it really 

] Davies, Geol. Mag-. 1880. ^ Lydekker, in Palaiontologiu Inclica, 1881-8G 

3 Lydekker, Fossil Vertebrata of India : Uecords Geol. Survey, India, vol. xx. 1887 
^ Op. eit. C;onii)t(!s rendus des SiSancos de I'Acad. d. Sci., Oct. 18HK. (An account of 
the fauna of Samos.) Atti della Societa Toscana di Scienze Natural!, I'roc -verb'di 
vol. V. 3 July, 1887, p. 272. (Letter addressed to Trof. Menegliini, in which' 
Dr. Major shows the difference between the mammalian remains of Samos and th o 
of Kos.) 
i Burckhardt, " Uber Aepyornis," Pal. Ahh., Jena, 1893, p. 21. 
'- Burckhardt, " Das Problem des antarktischen SchopfunMcentrums " &c Zool 
Jahrbiicher. 1902, p. 2(1. ' ' 

' Bull. Ac. Imp. Sc. St. Petersboura-, vol. xviii. 187.^. 


210 MR. F. E. BEDDARD ON [Mar. 3, 

is an egg of a Struthio has been proved by von Nathusius. Its 
geological age cannot certainly be determined because it was found 
floating in a river. Brandt supposes that it had been embedded 
in the bottom of the river and had been worked out by the water. 
The age of the sti'ata forming the bed of that river is supposed to 
be that of the strata in which von ISTordmann ^ discovered the 
mammalian remains in S. Russia (environs of Odessa), and therefore 
that of the breccia of Pikermi ; but it may be much younger, 
because Nordmann^ does not separate the Tertiary from the 
Pleistocene. In the paper above mentioned, he attributes the 
bone-beds of S. Russia to the "offenen Diluvium," and says that it 
has the same geological age as the bone-beds of the Val d'Ai-no. 
Therefore we have no evidence of the exact geological time during 
which a Struthio lived in South Russia, and the egg is in con- 
sequence of little importance in regai-d to this question. Moreover, 
it seems to me more than doubtful to assign this egg to the 
modern Ostrich, and "very likely to the species occurring at 
Samos," as is done in Nicholson and Lydekker's ' Manual of 
Palaeontology ' (p. 1228). 

5. On a new Genus and two new Species o£ Earthworms o£ 
the Family Eudrilidce, with some Notes upon other 
African 01igocha?ta. By Frank E. Beddard, M.A., 
F.R.S., F.Z.S. 

[Received March 3, 1903.] 

(Text-figures 35-38.) 

The first set of specimens referred to in the following descriptions 
form a part of the collection in the British Museum ; I am greatly 
indebted to the kindness of the Director for allowing me the 
opportunity of examining them. They belong to two species, 
both of which are undescribed. The first is a third species of the 
genus Stuhlmcmnia, which I call 

Stuhlmannia michaelseni, n. sp. 

I have examined so many individuals of this form of Stuhl- 
mannia, and the agreement between them is so close, that I have no 
hesitation in regarding it as a distinct species, which I name 
after the founder of the genus. Dr. Michaelsen ^. 

The worms were collected by Mr. S. L. Hinde in the Mt. Kenya 
district. There were thirty mature examples. 

The general appearance and proportions of this new species are 
quite the same as in ^S*. variabilis. The length reaches 100 mm. 
and the diameter 2-2*5 mm. The colour is a yellowish grey ; the 
clitellum is greyer. 

1 Nordmann, in ' Jubilteum semisaeculare Fisclieri de Waldheim,' Moscou, 1847. 

2 Nordmann, Palaontologie Siid-Russlands, 1858. 

3 JB. Hamb. wiss. Anst. vii. (1890), p. 24. 




Tlie setye ai'e strictly paired and the ventral pair of segment xvii. 
appear to be missing. The clitellum occupies segments xiv.-xvii. 

The median orifices of the spermatheca and of the spermiducal 
glands are very obvious, and frequently lie upon projecting papillae. 
The spermathecal poi'e opens far back upon the xiiith segment. 
The male pore opens far back upon the xviith segment, or even 
intersegmentally (xvii./xviii.). The penial settc can often be seen 
to protrude from it. Among the external characters, however, 
there is one which distinguishes the present species fi'om its allies, 
and that is the total absence of the penial pi'ocessof the body- wall. 
There was not the least ti'ace of this oi-gan obsei'vable in any of 
the individuals. It is true that this penis is not always to be 
seen in Stuhlmanida varioMlis ; but I do not think that it would 
be likely to be absent fi-om thii-ty individuals selected at i-andom ; 
and this absence is therefore one of the reasons which lead me to 
create a new species. 

Text-fiff. 35. 

Dissection of Stulilmannia michaclseni. 

xi, xii, segments flevi'u and twelve ; L.V., intcstino-tegumentav.v trunks ; o.d., oviducal 
apparatus ; Sp., spermathecal sac ; Sp.ffl., spermiducal glands. 

The accompanying draAving (text-fig. 35) illustrates a general 

212 MR. p. E. BEDDARD ON [Mar. 3, 

view of the viscera seen on cutting open the body-wall in the 
dorsal median line. No such figure of the anatomy of this genus 
has been published up to the present ; and it is convenient to 
show the relations of the different organs and their comparative 

There are points in the anatomy of the genus Stuhlmannia 
which are illustrated by that drawing, and to which attention does 
not appear to have been called. The last pair- of hearts lies in the 
eleventh segment, as in so many other Eudrilids, for instance in 
the genus Polytoreutus ; and the septum which should separate 
segments xiii. and xiv. is nearly missing. I am disposed to 
associate this latter fact with the presence of the ccelomic sacs 
svirrounding the gut, which may have been developed at the 
expense of the septum, as in Euch-ilus ', in which genus the 
corresponding septum is also much reduced. Another matter to 
which I would wish to direct attention, is the existence of intestino- 
tegumentary trunks, which have not yet been recorded in this 
genus. They are, of course, of wide occurrence among earth- 

A further point of some little interest is visible in the sketch 
exhibited herewith. In my ' Monograph of the Oligochteta,' '" I 
pointed out the existence of at least occasional asymmetry of the 
female reproductive organs shown in the presence of only a single 
receptaculum ovorum, that of the opposite side of the body being 
absent. In other specimens ^ I found precisely the same state of 
affairs. The species that I examined was, I believe, Stuhlmannia 

In the present species of StiMmannia there is exactly the same 
asymmetry, the I'eceptaculum being only present upon one side, 
and that the right. Or, to be more accurate, the receptaculum is 
possibly present also on the left side, but is quite rudimentary, 
and, I imagine, f unctionless. The " Eitrichterblase," as Mithaelsen 
has termed it, is present on the left side, and is simply a loop of the 
oviduct, the two sections of the tube running in close contact side 
by side ; just opposite to it is a small spherical projection. A 
lumen is present in this, which is the rudimentary funnel, but 
the lumen is no wider than that of the oviduct elsewhere, and 
there is no question of any free communication with the body- 
cavity outside ; thei-e was no break to be detected in the muscular 
wall of this projection, which perhaps should be regai'ded as funnel 
+ receptaculum. 

A series of sections through the rudimentary funnel and the 
adjoining parts of the oviduct confirms the appearances displayed 
in a preparation mounted in glycerine. The rudimentary funnel 
(or funnel + receptaculum) is but a slight protuberance, which is 
traversed up to its very end by a blindly ending branch of the 

1 Beddard, " On the Gonad Ducts and Nephridia of Eudrilus," P. Z. S. 1902, 
vol. ii. p. 89. 

2 Oxford, 1895, p. 602. 

3 " On some Earthworms from British East Africa, &c.," P. Z. S. 1901, vol. i. 
p. 355. 


descending limb of the loop, which constitutes the 
" Eitvichterblase " of Michuelsen. No communication between 
this slioi-t diverticulum of the oviduct and the general crelomic 
cavity could be detected. Evidently, however, it is the forerunner 
of the funnel which opens into the fully developed receptaculum, 
or, possibly, of both. It is noteworthy that the oviduct 
is divisible into two regions. The section lying nearest to the 
spermatlu'cal sac is of a wider calibre than that lying nearer to the 
external orifice. Furthermoi'e, the lumen of this wider section 
is a straight tu])e, while the lumen of the narrower section of 
the oviduct is sinuous in its coui'se within the muscular sheath. 
It is possible that the wide straight section is really to be looked 
upon as a greatly drawn-out funnel, drawn out by rea,son of the 
gi-owth of the spei'mathecal sac. 

The walls of the oviduct, as is the case with other Eudrilida?, 
are very stout and muscular ; and attention has ah'eady been 
called to the fact that part of tlie oviducal tube runs a, sinuous 
course, independent, therefoi'e, of its musculai- covering. These 
facts, coupled with the conditions obta,ining in EiulrilK.s ', where 
the oviduct runs for a considerable space actually within the 
septum, lead me to consider that the muscular wall of the oviduct 
in StiMmannia may be a, purely adventitious slieath, not belonging 
to the oviduct at all, but consisting chiefly of tlie otherwise 
missing septum between segments xiii./xiv. Its continuity with 
the receptaculum can at any rate be explained on this view, which 
is in any case not at variance with the other facts to which I have 
just called attention. 

The present species, in correlation, it is to be presumed, with 
the entire absence of a penial process, does not possess the median 
unpaired bui'sa pi'opulsoi'ia which chai'acteiizes *S'. variabilis. In 
the former point ^^'^''^^iWy, but not certainly, it agrees with 
S. gracilis. Di-. Michaelsen states, only as a possibility, since he 
only examined two specimens, the absence of a copulatoiy process 
in S. gracilis ; furthermore, a thickening of the integument of 
segment xv. appeared to him to be possibly intelligible as a not 
fidly developed penis. With I'egard to the presence or absence of 
a Inu'sa propulsoria he is silent. Stuhhmmnia michaelseni, however, 
caiuiot be confused with S. gracilis by reason of the characters 
of the penial setag. They ai-e provided at the end with a row of 
shaip denticles on either side as in aS'. variabilis. In short, I 
think that a case has been made out for the creation of a new 
species of this genus StiMmannia. 

Bettonia lagariensis, n. g, k sp. 

Of this apparently new genus and species I have examined but 
a single specimen, complete and sexually mature. It measures 
90 mm. by 4-5 mm. in breadth. The colour is bluish violet. It 

1 Bcddavd, " On tlie Gonad Ducts and Nqihiidia of Etidrilus," P. Z. 8. 1U02, 
vol. ii. p. 89. 

214 MR. F. E. BEDDAEB ON [Mar. 3, 

was collected by Mr. Stuart Betton, at Lagari, British Central 

The prostomium is continued by grooves over about half of the 
buccal segment. 

The setee are wider apart in the case of the ventral couples than 
in the case of the lateral. 

The limits of the clitellum are a little obscure ; it appears to 
embrace segments xiv.-xvii. inclusive. 

The chief, indeed practically the only, reason which leads me 
to separate this Eudrilid generically, is the condition of the repro- 
ductive apertures. The rule in the family is that the apertures 
of the sperm-dvict and of the spermathecse are unpaired and 
median in position. 

Text-fig. 36. 

V i 

Ventral surface of Bettonia lagariensis, ?, sperma thecal pores; <?, male pore. 

There are, however, a few exceptions, such as Eudrilus itself, 
and an apparently close ally of the present genus, viz. Eriiinoscolex. 
Bettonia offers a third arrangement of the reproductive pores. 




The spermathecal pores ai-e paiied, while the male poie is single 
and median (text-fig. 36, p. 214). 

The two former poies are upon the boundary line of segments 
xii./xiii. ; each is somewhat eye-sliaped in contour and corr&sponds 
exactly in position to the outermost of the vential couple of setae. 

Tlie single and median male pore lies on the boundaiy line of 
segments xvii./xviii. Jt is very large and conspicuous, with 
radially folded maigins, indicating, pei'haps, the possibility of the 
extrusion of the bursji propulsoria. 

The internal oi-gaiis, unfoi-tunately, were much softened, and 
therefore the female reproductive apparatus was rather difficult to 
deciphei'. The nature and i-elations of the other organs of the 
body were not so diflicult to asceitain. 

As in many, if not in all, Eudiilidse, the last pair of hearts 
occurs in segment xi. In front of these were four distinct pairs. 
The doi'sal vessel is single thi'oughout. I have not been able to 
study the alimentary system in detail, but I have ascei-tained 
that the present species is a member of the section Eudrilacea l)y 
virtue of the existence of an unpnired ventral median alimentary 
gland in segment x., which showed on a niicroscopical examination 
the usual laminate structure of these glands, not at all to be 
confounded, even in a badly preserved specimen like the present, 
with the totally difterent though corresponding organs in the 
other subfamily of the Eudrilidae. There are also a pair of these 
glands in segment xiii., of an oval form, which contain abundant 
crystals, such as are met with in other genera. 

Text-fig. 37. 

Tei-mination of the male efferent apparatus of Bettonia lagariensis. 
B.p., Bursa propulsoria ; Sp.ffl., spermiducal glands. 

The only conspicuous parts of the male generative system are 

216 MR. F. E. BEDDARD ON [Mar. 3, 

the sperm-sacs and the terminal apparatus, which opens on to the 
exterior in the xviith segment. The sperm-sacs are in segments 
xi. and xii., and occupy a considei-able space in those segments. 
They are simple, solid, sac-shaped structures, not racemose in form. 

The spermiducal glands are paired. Each of them (see text-fig. 
37, p. 215) is shortish and rather thick, tapering somewhat towards 
the tip. The last third of the gland is bent forward and lies above, 
parallel to and in contact with the anterior section of the gland. 
This arrangement occurred on both sides of the body. Each 
gland is sharply constricted at its opening into a large median 
bursa propulsoria lying below the ventral nerve-cord. This latter 
sac, however, presents obvious signs of having been produced by 
a fusion of two sacs ; for posteriorly it is completely double. It 
is into each of these posterior lobes that the spermiducal glands 

My description of the female appaiutus must unfoi'tunately be 
incomplete. The oi'gans, as already stated, are paired. The 
spermathecfe, near to where they open, have very thick musculai' 
walls ; and this i-egion at least is enveloped in a coelomic sac, as is 
the base of the spermatheca in the species of Pareudrilus (?), with 
which I deal later in the present communication (see below). 
How this ccelomic sac is otherwise related to the spermatheca 
and to the receptaculum ovorum I am unable to state. The latter 
organ presents no noteworthy peculiarities, and the ovidiict leads 
from it to the exterior, on the fourteenth segment. 

On a Sjyecies of Pareudrilus. 

I believe that a number of individuals belonging to this genus, 
which were collected by Mr. Crossland, may represent a new 
species. But I am unable to speak with absolute certainty on the 
matter, since the material was not in good order for investigation, 
and since the specimens of P. papillata examined by Michaelsen ^ 
were likewise much softened by evaporation of the alcohol ; if the 
worms upon which I report here are not referable to P. 2)apiUata, 
then the species is unqviestionably new. 

But whether the species be new or not, I have something to add 
to what is known about the structure of this genus. 

The dimensions of my specimens agree apparently with those 
given by Michaelsen for his sj)ecies. The length was some 100 mm. 
and the diameter about 3 mm. The dark purple colour, turning 
to yellow below and in the clitellar region, is like that of my 
Pareudrilus stagncdis. The setee are closely paired, and I observed 
a tendency in the neighbourhood of the genital pores for one seta 
of a pair to be lost or not developed. I do not refer to a mere 
dropping out ; on examining the cuticle, occasionally no pore was 
to be noticed in the place where such a pore (through which the 
seta is extruded) should be. The irregularity of this state of afi'airs 

' " Die Regenwurmer Ost-Afrikas," in Deutsch-Ost-Afrika, p. 11. 


does not incline me to describe in detail the cases observed, which 
were, moreovei-, not many. I can eoniirm Michaelsen's statement 
that there are no genital or any specially modified setfe in any 
region of the body. 

In the external sexual chai-acteis I find some little diflerences 
from the description of Michaelsen, and it is pai-tly on this account 
that I am dis[)osed to regard my specimens as belonging to a new 
species. The clitellum is perhaps a little more extensive, xiii.- 
xviii. or even xix. ; but that is a slight difference. I find that the 
pores of the paired spermathecie are situated definitely on the 
l)order-line of segments xiv./xv. and not upon xiv. They are, 
however, in the line of the ventral couple of setre. On the other 
hand, I agree with Michaelsen in placing the pores of the spermi- 
ducal glands upon segment xvii. just in front of the ventral setae 
of that segment. As to the location of the genital papillae, l_>r. 
Michaelsen and I have not found the same conditions in the 
specimens studied. In my specimens, the constant rule appears to 
be the pi-esence of two pairs of pajiilLe of which the most anterior 
are on segment xvii., just behind the ventral seta^ but on a line 
with the outermost setre of the couple ; on the following segment, 
the xviith, there is a precisely similar paii' also behind the ventral 
set;e, but in this case corresponding to the innermost seta of the 

The papilhe aie faiily conspicuous, and aie to be noted upon the 
cuticle when stripped off. The area is slightly laised and is 
studded with the mouths of large glands. Corresponding to the 
papillre inteinally, are glands to which I shall recur in describing 
the anatomy. These papillae were nevei- absent. 

I could discover no nephridial pores ; but I imagine that this 
species, like Fareudrihcs stagualis and some other Pareudrilacea, 
will turn out to possess a ramified system of nephridial end- 
tubes in the integiunent. In any case the nephridia, viewed 
inteinally, appear to be paired structures. As to the alimentary 
tract, I find a gizzard far foi-ward, in the sixth or seventh segment, 
and I have not been able to find calciferous glands. The last 
heart is in xi. The funnels of the sperm -ducts are in x., xi. The 
two paii's of sperm-sacs, in xi., xii., are tongue-shaped. 

As to the female generative system, my observations do not 
altogether agree with those of Michaelsen ; but it is no disparage- 
ment to that excellent and accurate obsei'ver to suggest that the 
condition of his specimens may possibly have led him into some 
slight error. Otherwise I must place my specimens not only in a 
new species, but in a new genus. The ovary I have not been able 
to find at all. This in an Eudrilid is not of course surprising; 
for as a i-ule the ovary disapjieai'S early, having transfei-red its cells 
to the egg-sac. On the other hand, while it is unlikely that I 
could have missed so conspicuous an object as the large o\'arian 
sac figured by Michaelsen, it is equally unlikely that he wa,s misled 
by bulging septa or other structures into stating the presence of 
such a sac communicating with the egg-sac. It seems to me 

218 MR. F. E. BEDDARD ON [Mar. 3, 

therefore to be possible that, after all, in spite of undoubted 
resemblances, Michaelsen and I have examined difterent species 
and indeed different genera. 

The female organs of the species which I have examined show 
a nvxmber of not uninteresting features. The two most salient 
parts of that system, which are visible on a dissection, are the two 
spermathecse and the egg-sacs. The spermathecse are sausage- 
shaped, and distinctly divisible into two regions. The proximal 
part, i. e. that nearest to the external orifice, is strongly muscular, 
and indeed is enveloped with stronger muscular bands than is the 
bursa propulsoria of the spermiducal gland. Its epithelium is 
perfectly continuous with the general epidermis of the body- wall, 
and it has every appearance of being formed as an ingrowth from 
the exterior. The distal region of the spermatheca has very thin 
muscular walls, much thinner than the walls of the glandular part 
of the spermiducal gland. 

The interior has an epithelium which is raised into folds. I 
cannot speak of the histological characters of the cells, as the 
material was not sufficiently good. There is a very close resem- 
blance, on a superficial view, between this spermatheca and the 
spermiducal glands. Indeed, on a cursory inspection, they 
might be taken for consecutive pairs of either spermathecse or 
spermiducal glands. The next most obvious part of the female 
reproductive system is a very large mushroom-shaped body closely 
adherent to the septum dividing segments xiii./xiv. This body is 
stalked, and appeai'ed, on dissection, too large to be identified 
with a receptaculum ovorum (or egg- sac). Nevertheless it is the 
egg-sac, and by virtue of its large size it appears to be precisely 
like the egg-sac of P. pajnllata described by Michaelsen. On a 
closer inspection, a fine tube, apparently leading from the stalk 
of the egg-sac to the muscular part of the spermatheca, was 
apparent ; this seems to correspond to the narrow tube (sg.) figured 
by Michaelsen. 

A series of longitudinal sections through the body showed more 
accurately the relations of these diverse organs to each other. I 
find that the spermatheca is entirely independent of the rest of 
the female apparatus, and that its cavity does not communicate 
with the narrow tube arising from the egg-sac. That narrow tube 
exists, as I have already mentioned ; but on reaching the base of 
the spermatheca, i. e. the muscular end portion, it dilates into a sac 
which entirely surrounds the muscular part of the spermatheca, 
but does not, so far as I could ascertain, open into it anywhere. 
The conditions, therefore, are those of such a genus as Hyperio- 
drilus or Heliodrilus, where a true spermatheca is invested by a 
ccelomic sac. Now, though the difference may appear to be slight, 
I am disposed to think that it is important, and that a sperma- 
theca which has no communication with the egg-conducting 
apparatus is essentially different from a spermatheca which has 
such a communication. It seems to me, for example, to be wrong to 
compare the spermathecal sac of Lybioclrilus or Stuhhnannia with 


the spermatheca of JTeliodrllus. For this i-eason I cannot agree 
to Mieluielseu's ])laciiig- of my genus Alvaiiia within tlie genns 
IL/pertndrilus. Tlie former Iims a, true spermatheca, homologous 
witli tliat of otlier non-Euih'ilid eartliworms, while the latter hi\« 
not. This also is the case with the genus or genera with which we 
are dealing n-ow. Pm'eiulrilus staynalis has a spermathecal sac 
which communicates with the c(elomic sacs envolving the ovary; 
" Pareudrilas " jmpillata has not. It is possible, therefore, that we 
should revive Michaelsen's Unyoria for the latter species. The 
stalk of the egg-sac, as might be expected, lodges the funnel of the 
o\iduct, or, to speak more accurately, the greater part of the funnel. 
The exact conditions obtaining ai'e the following. In a series of 
sections it may be seen that the nai'row tube communicating with 
the ccelomic sac siu'i'ounding the proximal end of the spei-matheca 
runs forwards and opens into the caxity of the xiiith segment by a 
wide oiitice ; its walls are therefore continuous with, and no doubt 
developed from, the septum bounding segment xiii. posterioi-ly. 
The uppei- " lip " of the ostium is covered by the cubical cells of 
the oviduct, which hei-e opens freely into the cavity of the xiiith 
segment. Further on in the series of sections, the mouth of the sac 
surrounding the spermatheca is closed, and the tube oj^ens into the 
egg-sac through a wide tube which is entii'ely lined by the cells of 
the oviducal funnel ; these are, of course, perfectly continuous with 
those cells wdiich lie in the xiiith segment. The part of the 
oviducal funnel which lies in the xivth segment appeals to be 
divided into two, and to open by as many mouths into the hu"-e 
egg-sac, which is so kidney-shaped as to be nearl}^ divided into two 
sacs; I cannot pretend to an accurate description of the funnel and 
its various foldings. It is clear, however, that the conditions which 
obtain ai'c those of the more typical Earthworms, where the funnel 
opens partly freely into the xiiith segment and is pai'tl}^ I'eflecterl 
so as to open within the egg-sac. Now, the simpler forms of 
Eudrilida?, such as the genus Eiidriloides, are distinguished by the 
fact that the ovary is unenclosed in any sac, and that the funnel 
of the oviduct opens piecisely as has been just described in Pareii- 
drilus jyapillcda. On the othei' hand, in the more complicated 
foi-ms, such as Stuhhnaniiia, the sacs containing the ovaries 
envelop also the oviduct-funnel and communicate with the 
spermathecal sac. The species which forms the subject of the 
present remarks is plainly intermediate between these two 
extremes ; and for that reason, as I think, deserves geneiic 
sepaiation fiom Pareridrilus. 

§ On the Sjiermatojihore. 
The spermatophores of the Eudrilida^ have not been much 
studied, and, so far as I am aware, are known only in the o-enera 
Stuhlmannia and Poly tor euttos, in which I ha^-e m}'self described 
them. Many, if not most, of the individuals of the present .species 
which I examined had a single spermatophore in both the 



[Mar. 3, 

As willTje seen from the accompanying drawing (text-fig. 38), 
the spermatophore consists of a globular swelling followed by a 
long thin tvxbe. The sperm, which appears blackish in glycerine 
pi-eparations, is limited to the spherical or nearly spherical recep- 
tacle at the distal end of the case. The spermatophore, as will be 

Text-fio-. 38. 

Spermatophore of Paretidrilns sp. 

also apparent from the figure referred to, is of an elegant form, 
not precisely corresponding to that of the spermatheca in which 
it lies. The swollen and globular receptacle of the sperm is not 
more than one-third of the length of the " stalk," which reaches 
down to the very mouth of the spermatheca. At its termination the 
Avails of the spermatophore project in a ring-like fashion ; a state of 
afi'airs exactly recalling the spermatophores of the Tubificidfe, and 
of the genus Stuhlmannia among the Eudrilidse. No doubt the 
shape is due to a moulding upon the walls of the spermatheca ; 
but the state of preservation of the specimens does not enable me 
to give details. The walls of the spermatophore appear to be firm 
and thick, and rather brittle in consistency. In teased prepara- 
tions the rupture of the stalk was invariably a clean fracture. 
The walls aie fibrous in appearance, and of the usual pale brown 
colour that is generally associated with chitinous membranes. The 
extremity of the tube, i. e. that which is nearest to the mouth of 
the spermatheca, is open ; the other end is quite blind. The 


shape of the whole sperniatophove is sufficiently ("hicidated by the 
drawing icfefied to. 

§ Note, oil, the CUlallniit (^'Alma stuidmainii, dial on a possibti/ 
new species of the genus Alma. 

I believe that a note by myself ' upon the clitellum and 
spermatophores of a West- African species of the genus Ahna is 
the first recoi'd of the extent of the clitelhun in that genus. Like 
other aquatic forms, Alma seems to be characteiised by a sea.sonal 
development of the clitellum ; and hitherto, with the exception 
just mentioned, no one appears to have seen or at least described 
this oigan in that Geoscolecid. In the species to which 1 have 
just referred, the clitellum was found to extend from segment xlv. 
to Ixxxv., a position which is quite unlike that found in any other 
Geoscolecid and present only in a few Lumbricids. This is an 
additional reason for associating the genus Alma more particularl}- 
with Criodrilus, as is done by Michaelsen ; for both these genera, 
though I'eferable to the family Geoscolecidne, have many points of 
kinship to the Lumbricida\ As, howevei', up to the present time, 
but one species of the genus Ali/ia has been descril)ed in the fully 
mature condition, it is possible that the position and extent of the 
clitellum characterising that species are not normal but exceptional 
in the genus. Theiefoi-e I do not hesitate to desciibe the con- 
ditions occurring in a second species of the genus, which I owe to 
the kindness of Mi-. Cyril Orossland, who collected specimens on the 
shore of Victoria ]Sr3'anza, among weeds cast up by the waves. 1 
have tw^o fully matuxe examples of a species which I believe to be 
identical with Dr. Michaelsen's Alma stuhlmanni. The dimensions, 
howevei-, are rathei' less ; only one of the two examples was quite 
intact — the other had lost the hinder end of the body; in the 
complete example, measurements showed a length of 120 mm. 
The other ujight have l)een slightly longer, as it was rather thicker. 
In both, the penial appendages were i-ather longer than those of the 
original specimens described by Michaelsen. I found them to be 
10 millimetres long ; in one example the two weie unequal in 
size, one penial appendage only mesisured 6 mm. The structure 
of these appendages is usually chaiucteristic of the species. The 
worms which T ha\e examined agreed in almost eveiy detail with 
the description of A. stuhlmanni as given by Michaelsen'-. 

I n»ay i-emark, howevei", that thei-e were only two setse at the 
free end of the penial process, and, indeed, one of these had 
dropped out. The two papilkv upon which these setje ai-e placed 
were paiily encircli^l by a, horseshoe-shaped region of specially 
tdandular epideiiuis, which was conspicuously marked out from 
the rest of the int(>gument covering that process. It is conceivably 
this region which secretes the spermatojihoi-e. In addition to this, 

1 "On theClitolluni ami S))iTiiiiitoi)limi'-^ utaii Aiiiidiil of tlic Gi'iius^/iwrt." Proc. 
Zool. Soc. 1901, vol. i. p. 21ii. 

- " Die Regeiiwiinner Ost-Afrikas. ' in Dinitseli-Ost-AtViku, 181)G, ]). k 


I fancy that the single papilla at the base of the penial process is 
rather nearer to the base than is figured by Michaelsen, and much 
nearer than in the West- African form, and, moreover, it appeared 
to me to be not symmetrical in its position. The clitellum was 
not seen by Michaelsen at all. In my specimens the clitellum 
occupied the same segments ; but, as in the species described by 
myself from Western Africa, that geneiative region was somewhat 
undefined in its beginning and ending. The greatest number of 
segments referable to the clitellum in the West- African species 
are from xlv.-lxxxv. ; but the clitellum was only fully developed 
upon segments xlvii.-lxxxii. Its general appearance was precisely 
like that of the species which I have just mentioned. In the 
present species — and the observations apply to more than one 
specimen, and are therefore all the more reliable as an expression 
of normal conditions — the clitellum was much shorter, only 
extending from segments xl.-lxxi. This rather leads me to the 
inference that the species with which I am concerned here is in 
reality difierent from that which I described fi'om McCarthy Island 
on the Gambia, and referred to A . stuhlmanni. The present worm 
is undoubtedly A. stuhlmanni; and it seems to be necessary, on 
account of the difference in the clitellum, to use another name for 
the West- African form. I would propose, thei-efore, to call the 
latter A. hudgetti. It is, however, clearly a very close ally of 
A. stuhlmanni. It is interesting to note that the species of this 
genus go more or less in couples. The East- African A. nilotica 
corresponds to my A . millsoni from West Africa, while A . stuhlmanni 
seems to be neaiest to the form which I propose to name J^. hudgetti. 
At present more information is wanted about A . emini ; but it 
appears to be formed rather after the plan of A. millsoni and 
A . nilotica. For in those species there are special setse on the penial 
processes, while in A . stuhlmanni and S. hudgetti there are setse of 
the same pattern as those on the body generally. 

March 17, 1903. 

G. A. BouLENGER, Esq., F.E.S., Vice-President, 
in the Chair. 

The Secretary exhibited, on behalf of Prof. Newton, F.K.S., 
three photographs of the White Rhinoceros {Rhinoceros simus), 
sent to him with the following letter by Mr. C. R. Saunders, 
C.M.G., Chief Magistrate and Civil Commissioner inZululand: — 

Eshowe, Zululand, 
^ „ 6th January, 1903. 

Dear Sir, 

I received a letter from you in August 1900, following 
on an account, written by me in ' The Field,' of an interview I had 
with White Rhinoceroses about that time. I did not answer 
your letter at the time, hoping I should be able before long to 
send you a photograph of the living animal. This, however, I 




liave not yet been iible to procure, although I visited the reserve 
in which they lived for this special purpose last winter. Their 
ti-aces were abundant, but my time was limited, and they could 
not be found. Thei-e are, I believe, about ten of these animals 
living in that reserve, and I do not despair of yet obtaining 
a iihotograph of them in life, in wdiich case I shall be pleased 
to send you a copy of it. 

Early last December two of the nninuds (lK)t]i ))ulls), one a very 
old one and the other not full-grown, str;iyed out of the reserve 
into one of the native locations and were killed. I obtained 
three photogriiphs of one of them, the old bull, t;iken l)y an 
amateur two or three days after it had died. I am forwarding by 
the same mail as this a. copy of each of these photographs, wliich 
you are welcome to, and which, 1 think, demonstrate the fact that 
they were taken from a specimen of the While Ithinoceros, although 
the carcass was a good deal distended. 

The killing of these two Rhinoceroses was most unfortunate. 
They suddenly appeared among some native kraals, and the men 
went out and attacked them with sjiears. The young one was 
killed outright; that of which the photograph was taken travelled 
a long distance after being wounded, aii<l was not found mitil 
some days had elapsed. Yours faithfidly, 

Alfred i\eivion, Esq. C. R. Saunders 

As tliese photogi'aphs weie piobably the only representations 
Text-%. 39. 

Kircnlly-killtd liliiiiorinis siiinis, aAn^i $. Dec. 1902. 

Proc. Zool. 8oc. — 1903, Vol. 1. No. XV. 15 


ever taken of this animal in the flesh, it seemed well worth while 
reproducing two of them (text-figs. 39, 40), 

Text-fig. 40. 

Recently-killed Rhinoceros' simus, adult $ . Dec. 1902. 

Mr. Oldfield Thomas exhibited the skin of a Chinese Monkey, 
which had been obtained from a hunter by Mr. Henry Brelich, 
and presented by him to the National Museum. It appeared to 
i-epresent a new species, the third known, of the remarkable genus 
Rhinopithecus, and was described as follows : — 

Rhinopithecus brelichi, sp. n. (Plate XXI.) 
Size very large, apparently larger than either R. roxellance or 
R. bieti ; for the skin, though that of a female, is as large as the 
male of either of the other species. Fur not abnormally elongated 
in any region, the longest being on the flanks, where the hairs 
may attain to about 90 mm. in length ; those of the back 50-60 mm., 
and those across the shoulders 70-80. General colour of back 
glossy slaty grey, the hairs grey to their roots, with shining tips. 
A pi'ominent oval white patch, 5 inches long by 2 broad, present 
in the middle line between the shoulders, its haiis white to their 
roots. Crown sufiiised with yellowish, its hairs yellow at base, 
whitening terminally, but with broad black tips ; haiis of cheeks 


STnit del.etlifh. 


Mon-ternBros . iirvp 


yellow with black tips ; nape between the yellowish crown and the 
white wither-jjatch pale brownish with black tips to the hairs ; 
neck both on sides and below blackish grey, the hairs dull whitish 
basally, with black ends. Eai'S white, markedly conti-asting with 
the head. Front of shoulders and inner aspect of foreai-ms deep 
yellow, which shades into whitish along the under aspect of the 
latter, and contrasts niai'kedly with the dark slaty of the outer 
side of the foi'earin, this colour darkening to black on the wrists. 
(The hands ai'e lost in the specimen, but are presumably black.) 
Hind limbs light greyish, more or less suffused with yellow behind 
and blackish in front, but the colour contrasts are not sharp and 
defined as tliey are in the other species. Belly uniformly grey 
(about grey No. 5 of Ridgway). Tail very long, conspicuously 
longer than in the othei- species, its hairs, which average about 
40 mm. in length, curiously curved on each side downwards and 
away from the centime line, along which there is an ii-i-egular 
parting ; in coloui' the tail is black thi'oughout except at the 
exti-eme tip, whei'e there is a small white pencil ; on each side of 
its base there is a small yellow patch, outside of which thei-e is a 
blackish line passing round across the anal region : but owing to 
the condition of the skin, the exact situation of these lines and 
patches is not quite certain. 

Appi'oximate dimensions of the type, measured on the skin, 
which has been made up from a flat native pelt : — Head and 
body 730 mm., tail 970 (with hairs 1040). 

Habitat. Mr. Brelich states that, " as far as I could gather, this 
monkey inhabits a. range of mountains known as the Yan Gin 
Shan Range, about 108° E., 29"" N., in the north of the province 
of Kwei-chow, Central China." 

Type. Female. B. M. No. Collected and presented 
by Henry Brelich, Esq. 

This magnificent Monkey, one of the lai-gest in the woi-ld apart 
from the anthi'opoids, is a very i-emarkable discovei-y, and one on 
which we may congratuLite Mi-. Bi-elich, who olitained and sent 
it to the Museum on the suggestion of Mr. Herbert Ingram, 
himself a frequent contributor to the National collections. 

As may be seen from the above description and from the figure 
(PI. XXI.), the differences between this monkey and its only near 
allies are so numerous as to render any detailed com2:)arison un- 
necessary. Good figui'es have been given of R. roxeUance by 
Milne-Edwards ^ and De Winton ^, and of R. bieti by Milne- 
Edwai-ds and Pousargues \ 

Mr. Oldfield Thomas also exhibited adult and young examples 
of a Bush Duiker, which had been sent to tlie British Museum 
by Mr. F. W. Isaac, from Eldoma Ravine, British East Afi"ica. 

1 Rcch. Mainin., Text, p. 233, Atl. pis. xxxvi. & xxxvii. (1874). • 

2 P. Z. S. 1899, p. 572, pi. xxxi. 

3 N. Artb. MuR. (3) x. p. 121, pis. 9-12 (1898). 



It appeared to belong to a new species allied to the Congolese 
Cephalophus iveynsi Thos., and was described as follows : — 

Cephalophus ignifeu, sp. n. 

Size medium. Fur close, fine and glossy, hairs of back about 
an inch in length. General colour of back bright rufous or bay 
(nearest to ochraceous-rufous of Ridgway), darkening forwards on 
the neck and shoulders to dull brownish. Forehead mixed rufous 
and black ; crown and occiput bright rufous like back, coronal 
tuft a deeper and more chestnut or vinaceous rufous. Hairs of 
occiput reversed upwards to the crest as usual, those of nape all 
directed backwards. Muzzle blackish ; lips and chin white ; ears 
dark bi-own behind, with white edges and inner surfaces. Throat 
rufous. Belly brown mesially, grading into rufous laterally. 
Inner side of forearms, inguinal region, and inner side of thighs 
white. Outer side of forearms and thighs I'ufous ; feet brown, 
darkening almost to black above the hoofs. Tail rufous above, 
white below, proximally, with a mixed brown and white terminal 
tuft. , 

Skull of normal proportions ; premaxillfe just reaching- nasals ; 
frontal convex as usual ; posterior palate vaiiable, that of the 
male cut out some way in front of the lateral notches, while in 
the female the median notch is posterior to the latei'al ones. 

Horns laid back just in the line of the face, those of an adult 
male 98 mm. long, with a basal diameter of 31 mm. ; the same 
dimensions in an immature female 49 and 20. 

The young specimen (skull 105 mm. long, the last milk jDremolar 
only just up) is nearly black all over, the coronal region and the 
posterior back alone being rufous. 

Dimensions of the type, an adult male, measured in skin : — 

Head and body 810 mm., hind foot, with hoof, (c.) 240, ear 81. 

Skull — basal length 162, greatest breadth 79*5, muzzle to orbit 
94, nasals 73x32; palate length 97 ; length of upper tooth-row 
53, of three upper premolars 23-4. 

Hab. Eldoma R,avine, British East Africa, alt. 7200 feet. 

T^/jue. Male. B.M. No. Collected and presented by 
F. W. Isaac, Esq. Three specimens. 

Native name " Meindet." 

This Duiker is allied to C. weynsi of the Eastern Congo and 
C. johnstoni of Toro by its colour and general characters, but 
difiers from them by the hairs of its nape being all directed back- 
wards in the usual way. G. harveyi of Kilima-njaro, also a 
member of this group, has a blackish forehead and no chestnut 
coronal tuft. 

The following papers were read :- 


1. Observations and Experiments on Japanese Lon<j;-taileJ 
Fowls. By J. T. Cunningham, M.A., F.Z.S. 

[Received January 1st, 1903.] 

(Text-figures 41 ct 42.) 

The enormous lengtli of the tail-t'eathei's in Japanese " Long- 
tailed Fowls " has been known to zoologists in this country for many 
years from the stutied specimens in the hall of the Xatui'al History 
Museum in London. These specimens were figured by the late 
G.J. Romanes in his 'Darwin and After Darwin,' Parti., puljlished 
in 1892. The figui'es represent fairly accurately two males of the 
breed, but the females are not represented at all, unless by indistinct 
figures in the backgi'ound ; and the accessories, instead of being 
di-awn from those actually present in the ^Museum case, are entirely 
imaginary, showing the birds in a state of perfect freedom in open 
country, a state which they never enjoy in Japan. Romanes's 
figures are among a lai-ge number given as " typical proofs of the 
eliicacy of artificial selection " ; and it is the object of the present 
paper to show that this involves an assumption, with regard to 
this particular breed, wdiich is by no means justified by the facts. 

The tails of the male specimens in the National Museum are 
6 to 9 feet in length, that is to say, the longest feathei's are of 
that length. I have endeavoured, but without success, to obtain 
information concerning the histoiy of these specimens. I l)elieve 
they were reared in Japan, and they were probably sent fi-om that 
country after death, either as skins or as stufied specimens. 
There are some female specimens also, but in these the tails are 
scarcely longer than in the liens of ordinary bi'eeds. 

The breed has been known to poultry-fanciers for a considerable 
time, and the following is the account of it given in ' The Book of 
Poultry' by Lewis \Yright, published in 1885 : — 

" About the year 1878 there appeared in CTermany, and a year or 
two later in England, fowls impoi-ted from Japan, wdiose principal 
peculiai'ity consisted in an immense length of tail and hackle- 
feathei'S. Some of these were exhibited as Yokohamas ; others, 
said to be superior iii these points, were called Phoenix fowls. 
The tails of these specimens averaged about a yard in length, and 
the genei'al appeai-jince was not only that of a Game-fowd, but all 
the colours were Game coloui'S, Whites, Piles, Duckwings, and 
later a few Black- Reds. The long plumage Avas, however, 
unique, and a fair idea of it may be gathei'ed from the illus- 

"Correspondence in the poultry journals brought out the fact 
that such birds had been occasionally exhibited as Japanese game 
.so far back as about 1872. But it further appeared that in the 
Japanese Great National Museum at Tokio there were pi-eserved 
two specimens, of an allied race in which the tail-feathei'S measure 

228 MR. J. T. CUNNINGHAM ON [Mar. 17, 

13| feet and 17 feet respectively, and a feather lias been actually 
sent to France which measures 2 metres 85 centims. in length 
(say 9| feet). In 1884 Mr. Gerald Waller, of Twywell, imported a 
pen of these biixls ; and from his statements we gather that they 
are known in Japan as Shinowaratao, Shirifuzi, or Sakawatao 
fowls and various other names. He says the very long-tailed 
ones are kept in high, narrow cages, always sitting on a perch 
covered with straw- rope, with no room to turn or get down, hut 
with a food- and water-tin at each end of the perch. Three times 
daily they are lifted down for a few minutes' exercise, their tails 
being carefully rolled up in paper cases to keep them from injury. 
The Japanese state that a tail has been measured 23 feet in length, 
and that the birds on\j moult the tail once in three years. This 
last statement is highly interesting. It is obvious that if a tail 
23 feet long were grown in one yeai-, it must be at the rate 
of nearly three-quarters of an inch per day ; and though 
Madame Bodinus states that she could see the tails grow daily, 
it is difficult to realise this ; but experience will soon decide the 
point. The birds which have reached Europe have never yet 
exceeded 5 or 6 feet in length of feather, which is not beyond the 
possibility of a single season, though it appears of an enormous 
length. The saddle-hackles of Mr. Waller's birds are 16 inches 
in length ; but it is manifest that such enormous feathers as 
reported from Japan covild never be preserved under the ordinary 
conditions of an English poultry-yard. The feathers are not only 
long but extremely nai'row and flexible, trailing low after the 

Mr. Wright does not mention the comb, but the illustration 
which he gives represents the male bird with what is called a pea- 
comb of small size and with small wattles, whereas the specimens in 
the National Museum have single vei'tical serrated combs and large 
wattles. The pea-comb is a rounded mass, with small rounded 
tubercles projecting from it. 

Mr. Frank Rice, of Acton, Sufiblk, who breeds Yokohamas, 
gives in his circular the same illustration which appears in 
Mr. Wright's book, and which therefore is certainly not a new 
figure from a living specimen. But in his description he states 
that the head should be neat and small, with evenly-set pea-comb. 
It would thus appeal' that the long-tailed fowls coiiiiDrise varieties 
which differ in comb as well as in colour, though they seem to be 
all similar in the excessive growth of the tail, and probably are 
all grown in Japan under the same artificial treatment. 

The principal purpose of this paper is to discuss the causes by 
which the elongation or excessive gi'owth of the tail has been 
produced. About two years ago, Pi'of . Lankester, in a letter to 
' Nature,' refeired to the specimens in the Museum of which he is 
Director as " a magnificent sport," comparing their exceptional 
character to what is called genius in human beings. On the 
other hand, in the ' Dictionary of Birds ' by Newton and Gadow, 
article " Feather," the length of the tail-feathers is attributed to 


continuous growth, and it is stated that the moult is checked or 
prevented by some means unknown to Europeans. We have seen 
that in Wright's book the statement is cited that the feathers are 
only moulted once in three years, but this might be a congenital 
peculiarity, whereas the expression used in the ' Dictionary of 
Birds ' implies that the moult is prevented by artificial treatment. 

Yery definite statements on the question are made in a paper 
by INIr. Basil Hall Chamberlain (" Note on a Long-tailed Breed 
of Fowds in Tosa," Tiuns. Asiatic Soc. Japan, vol. xxvii. 1900), 
for my knowledge of which I have to thank^ Mr. Frank Finn. 
Mr. Chamberlain made enquiries on the subject in the country 
from which the breed is obtained, and where presumal)ly it 
originated, but nevertheless his statements are based on the 
assertions of l)reeilers and not on any investigations of his own. 
The following is a summary of his paper : — 

The origin of the Ijreed is not knowai, Init it is believed to be at 
least a hundred years old. It has been produced simply by 
selection of the best specimens ; one highly-prized variety, the 
Haku, was produced in this way within the last few^ yeai-s. 

The proper general name for the Long-tailed fowds is Shino- 
wara-to, derived from the village of Shinowara in the province of 
Tosa, east of Kochi the capital. Some are still bred in that place, 
but most now in Kochi itself, whence the majority are exported 
to Kobe and some of the finest to Tokio, but the very finest are 
I'etained by the producers. 

The following varieties were described to Mr. Chamberlani : — 
Shira-fuji : white head- and body-feathers, tail black as in the 
other vai-ieties. He saw one specimen of this two years old, and 
measured its tail-feathers, which were 7^ feet long. Another 
specimen, fourteen months old, had tail-feathers 4 feet long. 
Others have black bodies. Another variety is the Haku, white 
all over with yello\v legs ; another, Totenko, has red neck- and 
body-feathers ; another, Dokiri, has reddish colour mixed with the 
white of the body. All these except the Haku have black tail- 
feathers. , 1 . 1 

As great a length as 18 feet has been reached m the tail- 
feathers, but even 12 feet is a rarity. From 7 or 8 to 11 feet is 
the usual length. The feathers grow about 4 inches a month, 
and continue "to grow while the Inrd lives, which may be eight or 
nine years. The beautiful body-feathers growdng from the 
shoulders reach a length of 4 feet. (iTo^e.— This evidently refers 
to the saddle-hackleSjVhich grow^ not from the shoulders but from 
the top of the rump.) 

Some of these saddle-hackles may fall ofl" in moulting, but the 
tail-feathers never do so. He saw the birds in Octol)er 1898 
when moulting, and only tlie ordinar}' feathers were gone or 
going, not the long ones. 

He also saw the hen, which was a handsome Inrd, distantly 
reminding one of a hen-pheasant, with fawn-colouretl breast and 
wdiite quill to the delicately-coloured feathers of the back. She, 

230 MR. J. T. CUNNINGHAM ON [Mar. 17, 

too, has longer tail-feathers than an ordinary hen, sometimes as 
long as 8 inches. One, or at most, two hens are allowed to each 
breeding- cock. The latter's tail-feathers are cut to allow of his 
walking about freely. He lives a little longer than the others 
which must be kept shut up ; but all are hardy, bearing both heat 
and cold. The ordinary number of long tail-feathers is 15 or 16, 
some cocks have as many as 24. 

The tail-feathers must not be wound up, as people ignorantly do 
away from Kochi, but must always be allowed to hang free ; for 
which reason the cocks are kept in high, nari'ow cages, quite dark 
except at the top, for light at the bottom would attract them. 
When the tail-feathers become too long and touch the ground in 
the cage, a bamboo is put a little way back so as to form an arch 
and make more distance. The birds sit all day on a flat perch 
3 inches wide, and are only taken out once in two or thi'ee days 
and allowed to walk about for half an hour or so, a man holding 
the tail all the while to prevent its getting torn or soiled. 

The high, narroAv cages may be made of any wood ; they are 
6^ feet high, 3 feet deep, and 6 inches wide. The wonderful 
feathers both on tail and body come from quills much stouter than 
any possessed by ordinary fowls. 

The price in Kochi was 15 dollars for a cock with tail under 
10 feet, 25 dollars over that length. 

There is absolutely no artificial method of making the feathers 
grow. All is done by selection. Any failure is due to not having 
a hen or parents of the proper breed. Also one must know how 
to treat the birds. 

At Kobe in November 1898 Mr, Chamberlain saw three speci- 
mens, one with tail-feathers 13| feet long. He also saw a splendid 
Avhite tail 10| feet long, which had been pulled out from a white 
bird owing to its falling off its perch and fluttering about. The bird 
was five years old, and the feathers were growing again. The fancier 
said that the feathers in young birds grow about 4 inches a month, 
in older birds more, up to 7 inches a month. 

Two photographs are given with the paper, but no reference to 
them is made in the text. The tails are very long, but there are 
no long feathers from the shoulders, only tail- and saddle- or 
rump- hackles. 

It is evident that Mr. Chamberlin, although his observations are 
of considerable value, was not an experienced naturalist, and that 
he is simply reporting what he was told. He says nothing about 
the combs of the fowls he saw. With regard to his assertions 
about selection as the sole means by which the breed has been 
produced, it is to be noted that he is evidently referring in the 
case of the Haku chiefly or entirely to colour. The Haku is a 
white variety, and Mr. Rice, above mentioned, also has a strain 
of this colour. Nearly all domesticated birds and animals vary in 
colour, and nothing is easier than to separate a white variety in 
fowls, horses, pigeons, dogs, &c. These fowls, like others, vary 
in colour ; and the question before us now is not the separation of 


colour varieties, but the excessive length of the tail which occurs 
in all the vaiieties. Mr. Chamberlain states that a specimen 
fourteen months old had a tail 4 feet long. At the rate of 
4 inches a month, this would mean a period of growth of twelve 
months, and therefore the chickens must have begun to acquire 
their permanent feathers at two months of age, and the fejxthers 
must have grown without intermission afterwai'ds. This is 
pei-haps not impossible, but accoi'ding to my observations, recorded 
below, it is very improb;iiile. It may further be pointed out that 
the statement " there is absolutely no artificial method of making 
the feathers grow," is ditiicult to reconcile w- ith that which follo\vs 
it, " also one must know how to treat the binls,"' unless proof is 
offered that the artificial treatment has no efl'ect on the growth of 
the feathei's. 

The birds in my possession, which have formed the subject of 
my own observations and experiments, were descended fi-om a 
pair which were imported direct from Japan by ^\v. John Sparks, 
of London, and purchased by ]\Irs. J. C. Williams, of Caerhayes 
Castle, Cornwall. Before proceeding to my own observations, I 
wish to discuss the question Avhether the moult occui'S in the 
specimens kept in this country, and also some evidence I have 
obtained as to specific treatment of the feathei'S. We have seen 
in j\Ii-. Wi-ight's account that the feathers of bii'ds in this countiy 
had not been known to exceed 5 or 6 feet in length. This 
does not thi'ow much light on the question of the moulting, 
because it may be that, the bii'ds not being kept on perches as in 
Japan, the feathei'S get bi'oken when they have reached a, con- 
siderable length. I enquii'ed of j\Ii'. Rice what his expeiience 
was, and he replied that his bii-ds usually moulted their tail- 
feathei'S each year, but he had had some cocks omit this operation 
in their second yeai-. He said that his cockerels at eight months 
of age had tails from 2 ft. 6 in. to 3 ft. 6 in. in length, and that 
he had one cock, three years old, that had a tail 5| feet long. 

Mrs. J. C. Williams was also kind enough to answer my 
enquiries. She informed me that the original male bird which 
she obtained from Mr. Sparks had a tail just 5 feet long. At first 
its plumage suffered from the change of climate, l)ut in 1901 its 
tail Avas considerably better than it had been. (The length at 
this time was not stated.) She had a young bird whose tail had 
measured 4| feet, but he broke it in getting about. The young 
birds usually moulted about a yeai' aftei' hatching. This must have 
been the first moult of the adult plumage, which begins to appear 
in the autumn aftei' hatching. In reply to a request for fuither 
particulars about the moult, ]\rrs. Williams stated the birds do 
certainly cast their long tail-feathers. The bird that had some 
feathers 44 feet long was then (INIarch 1902) nearly three years 
old, and she thought it Avas at the second moult that a feather of 
that length was measured. 

There is evidence here that, when left to themselves, the long 
tail-feathers are moulted in the ordinary way, at any rate for two 

232 MR. J. T. CUNNINGHAM ON [Mar. 17, 

successive seasons after the first year in which the birds are 

It will be seen that the absence of the moult, or the occurrence 
of continuous growth, if true for the birds in Japan, is not true 
for those reared in this countiy ; and thus we have i-eason to 
doubt that uninteiTupted or continuous growth of the tail-feathers 
is a fixed congenital peculiarity in the breed. 

Mr. John Sparks in May 1901 supplied me with the following- 
information concerning the method of treatment applied to the 
birds in Japan : — ^" In ordei- to ensure veiy great length of tail, 
the cocks ought to be kept on a perch as much as possible after 
they are six months old ; and the tail-feathers should be pulled 
gently every morning, grasping the centre bone-like part firmly 
with the fingei' and thumb and pressing steadily downwards 
towards the tip, each feather being done several times. This 
softens the quill and causes it to lengthen. The birds do not 
moult the tail-feathers, but if one or more come out others 
immediately grow in their places. 

" The Japs themselves, those who take great pride in their birds, 
always roll the long feathers up like a lady rolls up her hair, and 
tie them, whenever the birds are let off their perches to walk 
about, which is about twice a day for an hour at a time. 

" I have often seen them thus treated in Japan, and those 
which Mrs. Williams and the Hon. W. Rothschild had from me 
were so treated on the voyage by the man in charge of them, and 
I sent them down to St. Austell in their regular perch-cages." 

Thei'e is here a detail in the treatment of the feathers which, 
so far as I can discover, has never been mentioned in any 
published account of the matter ; and my own experiments, which 
I now proceed to describe, so far as they have yet gone, tend to 
show that this mechanical treatment of the feathers is the whole 
secret of the mystery. 

My own Experiments. 

In May 1901, I received from Mr. John Sparks twelve eggs, 
laid by the fowls of this breed in the possession of Mrs. J. G. 
Williams. I afterwards ascertained that some of these were the 
produce of the pair originally imjDorted from Japan, others were 
from the offspring of this pair, but all the eggs were of perfectly 
pure breed with no cross or admixture whatever. The eggs were 
incubated by an English hen of mixed breed, and on June 13th 
ten healthy chicks were hatched. Of the other two eggs, one 
contained a dead chick, the other was addled or not fertile. This 
shows that the eggs were remarkaljly fertile and of great vitality. 

The chicks were small, the eggs being not much more than half 
the size of ordinaxy table-eggs. Their colour was fawn, with a 
broad dark brown stripe down the middle of the back, a narrower 
stripe on each side of this, and a thin stripe of the same colour 
rimning from the outer corner of the eye. They were very 


pretty, very active, and very lieixlth}-, and tlii'ove well on ordinary 
food, coiLsi.sting of oatmeal, chopped meat and vegetables, and 
mixed grain of vai-ious kinds. One of the chicks was accidentally 
killed when I turned the hen out of the nest, having got beneath 
its mothei-'s feet as she was scratching the earth. I examined 
this specimen, and found that the primaries and secondaries of 
the wings were pi-esent as short black quills with a little down at 
the tip, l)ut that there was no trace of tail-feathers or tail-coverts, 
nothing but down over the rest of the body. The comlj was 
visible at the l)ack of the beak as a slight yellow ridge with six 
teeth. The toes were four in number ; the skin on the legs 
was yellow. 

Jane 23;yZ. Age 10 days. — Another chick had been lost u[) to 
this date, having escaped and probably fallen a ^•ictim to a cat. 
The feathers of the wings now i-eached nearly to the end of the 
body, and wei-e chequered in colour, l^eing mai-ked ti'ansversely 
with dark colour and gi-ey alternately. The tail-feathers, i. e. 
rectrices, had begun to sprout in some of the chicks, elsewhere 
thei'e was still only down. The combs ajipeared no larger. 

Jahj Gth. Age 23 dai/s. — The chicks were now half-fledged : 
anothei- had been lost by escaping, so that only seven survived. 
Feathei's wei'e growing on the shouUlers and sides of the bi'east, 
but the head, back, and fi'ont of the breast were still downy, with 
the original markings. Tail-coverts also appearing. Feathers, 
except tails, all barred with dai'k and grey, so that the general 
appearance was speckled and veiy inconspicuous against the soil. 
The tail-feathei'S were dark, near!}' black, Ijut not steel blue-black 
like those of the adult. It was interesting to see the chicks scatter 
in all tlii-ections, and then ci'ouch down whenever the hen uttered 
her special warning ciy, as she did often when she heard a jackilaw 
ci-oak. In four of the chicks the tails were more developed than 
in the rest ; in these also the combs were beginning to grow 
higher and to get red. These foui' pi-oved to be cocks, so that the 
sexual difference begins to show itself at this eai-ly age. 

Jiobj 28th. Age 1 month 15 dai/s. — ^The four cocks showed 
their sexual charactei's a little more distinctly, the wattles and 
ear-lobes being indicated by a tinge of I'ed. In the three hens 
the combs had not l)egun to enlai-ge and showed no i-ed. 

One of the cocks was darker than all the other chicks, and had 
a slender tail, not very long and l)ent downwai-ds. The hens were 
light -coloured, with white bi'easts, their tails being as long as those 
of some of the cocks. Two of the cocks had dai-k breasts. 

Aiog. 4ith. Age 1 vwnth 3 toeeks. — Chicks now fully fledged. 
In the four cocks reddish tints were appearing in the feathers of 
the back and wings, while in the hens only neutral tints were 
present. The combs in the cocks were a little more developed, in 
the hens not developed. The tnil- coverts in the cocks were growing, 
but not longer than the rectrices, and the sickle-feathers not 

The red feathers mentioned' above are the beginning of the 

234 ME. J. T. CUNNINGHAM ON [Mar. 17, 

adult plumage. There is at least one complete change of feathers, 
from the chicken plumage to the adult, both in cocks and hens. 
Whether the chicken-feathers are changed before this I do not 
know, but there were indications that the wing- feathers at least, 
in the chicks, had been changed before the assumption of the adult 

Aug. 12th. Age 2 months. — I examined one of the cocks, and 
could distinguish two steel-blue sickle-feathers on each side, not 
longer than the rectrices and rather narrow. These were the first 
of the adult tail-feathers to appear, succeeding others of dark, 
lusti'eless, bi'ownish black. 

Aug. 20th. Age 2 months 1 iceek. — I counted the tail-feathers 
in some of the chicks. In one of the hens I found 14, or 7 pairs, 
of rectrices. In one of the cocks, the dark one previously men- 
tioned, the saddle-hackles were almost black with a grey stripe 
down the centre, instead of reddish. In this bird I found also 
seven pairs of rectrices, and only four pairs of sickle-feathers 
which were neuti-al bi'own. The other three cocks could also now 
be individually distinguished, and I give the peculiarities to show 
the amount of variation : — 

A. The largest : more advanced in plumage than the rest, more 

red on the back, breast nearly all black except at the sides ; 
a kink in the comb. 

B. Similar to A, but not quite so far advanced. 

C. Similar to A and B, but lighter in tone ; back rather 

yellowish than red. 

D. The dark bird above mentioned. 

Au,g. 2Ath. Age 2 months 11 days. — Examined Cock A and 
found a number of steel-blue sickle-feathers, or tail-coverts, all 
with very long horny sheaths at the base indicating vigorous 
growth. In Cock B similar feathers were only just beginning to 

Aug. 2&th. — In Cocks C and D very few new sickle-feathers 
showing. In B the rectrices were being replaced. 

Sept. Qth. Age 2 months 3 loeehs. — In Cock A all the chicken- 
rectrices had been moulted, and the new steel-blue ones were 
growing out with long horny sheaths at the bases. In all the 
cocks the sptirs had begun to show as very slight blunt knobs. 
ISTone of the cocks had crowed yet. 

Sept. 10th. — Examined one of the hens. Saw several tail- 
coverts growing with long sheaths but no new rectrices. 

In Cock D, the dark one, the two central rectrices were long, 
slender, and curved, and had sheaths at the base, also sheaths to 
some of the other rectiices. I thought at the time that these 
central rectrices had not replaced others in the chicken plumage, 
but grown continuously. I afterwards concluded that they had 
moulted earlier than in the other cocks. 

In Cock A the longest tail- covert measured 19 cm. or about 
7 1 inches. 


tie-pt. 1 9i/i. 2i(je 3 months 1 loeek. — Condition of plumage : — 

Cock A. Breast, thighs, and l)elly all black. Xeck-hackles 
very light grey, with thin dark stripe down the centi-e of 
each feathei'. Back with some steel-blue feathers behind 
the hackles, the rest red. Saddle-hackles developing, 
yellow, long and thin. Nearly all the rectrices and tail- 
covei'ts growing with long sheaths, colour steel-blue. 

Cock B. Nearly the same as A, but not quite so far advanced. 

Cock C. Much less advanced, white on sides of breast, speckled 
brown feathers mixed with red of back, only the outer- 
most rectrices shomng sheaths. 

Cock D. Very little white on breast, hackles of neck steel-blue, 
back black with a little red at tips of feathers. Saddle 
whitish. Rectrices 7 pairs, only the central and outer- paii-s with sheaths, the rest appai-ently not yet 

Hens. Breasts almost white, the sides of In-east bufi"; neck- 
hackles dai-k, with whitish stiipes down centres of f eathei'S. 
Back and tail greyish bi-own, speckled, i. e. with white 
quills. Two hens with black heads, one Avith head lighter, 
grey and speckled. 

I now decided to stroke and pull the tail-feathers in Cock B and 
to leave Cock A untouched, to see if any difference would result. 
These two, as I have said, were closely similar except that A was the 
larger', finei- biixl and slightly more advanced in development. I 
chose, thei-efore, the one which w'as congenitally inferioi-, so that 
if an}'- superiority in growth of feather appeai-ed in it, it could 
only he due to the artificial treatment. In both thei-e wei'e seven 
pairs of rectrices, the central pair slender and curved, the rest 
broad and straight. The outermost paii- had not yet been 
moulted. I fixed up a sort of cage with a round pei-ch at the 
bottom, and put Cock B into it while I stroked his feathers, but 
did not keep him in it. 

Oct. \st. — Found that the hens had shed most of their rectrices 
and were producing new feathers, as well as new tail-coverts, liut 
without change of coloiu-. 

Oct. 1th. — The ui^per tail-covert in Cock A measured 25 cm. or 
about 10 inches, not including the basal sheath. 

At this time I changed m}' r-esidence, and the fowls were 
installed in a place divided into two runs ; into one I put Cocks B, 
C, aird D, in the other Cock A with the three hens. 

On Oct. 12th Cock A crowed for the first time, another proof 
that he was a little more precocious than the others. 

Oct. \^th. Age 4 months 3 c?«..ys.— Longest feather, a tail- 
covert on left side, in Cock B 27 cm., about the same as the 
longest in Cock A. 

Oct. '27th. Age 4 months 2 loeeks. — Tried tying the cocks by 
one leg on ordinaiy perches about 3 feet from the ground, and 
thus was able to measure the feathers better. 

236 MR, J. T. CUNNINGHAM ON [Mar. 17, 

Cock A. Longest feather 13 inches from skin. 

Cock B. The same. But there were 7nore long feathers in 
Cock A, the outer ones being more nearly the length of the 

As I found I had not time to pay attention to more than two 
cocks, and as, moreover, they soon began to fight, I got rid of 
Cocks C and D ; and what I have to say hei'eafter refers only to 
Cocks A and B, which were undoubtedly the best of the four, and 
had the most beautiful coloui's. I began tying these two cocks 
on ordinary perches in the daytime by means of a piece of tape 
tied to one leg and round the perch. 

Bee. 1st. Age 5 tnonths 18 days. — Longest feather in Cock A 
18 inches, in Cock B the same. This shows a growth of 5 inches 
in five weeks, oi- 1 inch per week. Pei'haps the feather in Cock A 
was really a little longer, as neai'ly all the feathers in this bird 
had lost their tips by friction against the ground. There is thus 
no evidence up to this time that pulling the feathers in Cock B 
has increased the rate of growth. 

Dec. 8ih. — I had been stroking the feathers of Cock B regularly 
at morning and evening. Usually I tied the birds on the perches 
at night, and generally tied the feathers up in paper when the 
birds were free. I used tissue-paper, and rolled the tail up trans- 
versely, fastening it with tape. I tried a rough naiTOw cage with 
wooden bars foi' Cock B, but the bird turned round and escaped 
through the opening behind the perch. 

I noticed now that one of the long feathers in Cock B seemed 
to have stopped growing, the sheath having peeled ofi" and left a 
dry stalk. This was one of the outer tail-coverts, but the rest 
were still growing vigorously. 

Dec. 27th. — As my first attempt at a cage did not succeed, and 
T was unwilling to confine the birds so completely as the Japanese 
do, I made a. perch of about 9 inches in length supported on two 
uprights. This apparatus was movable, not fixed to the ground. 
I used this for Cock B, tying him to it by a piece of tape fastened 
round one leg. This answered veiy well, although he once upset 
the perch and slightly wounded his head. No permanent injury 
resulted from this accident. 

On this date, when I was sti'oking the feathers, one of the 
smaller tail-coverts on the right side came out of the socket, with 
the epidermic sheath attached to it. This feather was 14 inches 
long. This seemed to indicate that the eflfect of stroking the 
feathei's was, as suggested by a correspondent of ' Nature,' to pull 
the gi-owing feathers out, instead of increasing their growth ; but 
my experiments were not finished yet. 

Dec. Slst. Age 6 months 18 days. — Measured the feathers as 
accui'ately as possible on a flat wooden measure. 

Cock A longest two feathers, the central feathers of the tail, 
22 inches. 

Cock B longest feather scarcely 22 inches, also a central 


It seemed, therefore, that the gi'owth of the feathers in Cock B 
was barely keeping pixce with that in Cock A, although only in 
the former were the feathers pulled or stroked. In Cock A the 
longest feathei's Avere the two central i-ecti'ices ; in B only one of 
the central rectrices was as long, the adult feathers having mostly 
staited earlier in A than in B. 

Up to this time Cock A had been with the three hens, and B 
with the other cocks until I had got rid of these. 

1902, Jan. 8th. — On this date one of the hens laid the first egg. 
It was small and of light brown colour, like those the birds were 
hatched from. 

The hens have a low seii'ated comb and scarcely any wattles, 
only red skin about the cheeks. 

Ja7i. I2th. Age 7 months. — In Cock A the 2nd pair of rectrices, 
counting from the middle, seemed to be coming to the end of their 
growth, the sheaths diying up and the quills forming. This pair 
wei-e not so long as the corresponding paii" in Cock B which 
had been pulled. The five outer pairs of rectrices had ceased 
growing in both cocks. These were bi-oad and stiff and only about 
6 or 7 inches long, in fact like the i-ectiices of ordinaiy fowls. 
All the hens were now laying. One of the hens was put with 
Cock B, the other two Avith A. 

Feb. Ath. Age 7 months 22 dai/s. — 

Cock A, longest feather 25i in. = 64 cm. 

Cock B „ „ 25| in. = 65-7cm. 

It is thus evident that the gi-OA\i;h in B had been greater than 
in A. The feathei'S in B wei-e pulled once or twice a day ^\•hen- 
ever I was at home, and B was tied on the pei-ch with his feathers 
loose at night, while A was usually left free with his tail I'olled 
up in papei-. The sheaths of the centi-al i-ectiices, which were 
the longest feathei-s in both birds, wei'e in Cock A shoi-ter, and 
seemed as though gi'owth were about to cease. 

It is to be noted hei-e that so long as the feather is in full 
growth, the horny sheath which surrounds the base is milky 
white, or bluish white, and soft, while when growth ceases the 
sheath dries up and appears black because the feather is seen 
through it ; then it scales off, and leaves the quill of the feather 
bai'e down to the surface of the skin. Tlie quill at the base of a 
full-growai feather is only foi-med at the end of growth, and most 
of it is contained in the skin-socket. Thus I leai-ned to I'ecognise 
the approaching cessation of growth and formation of the (piill by 
the appearance of the horny sheath. 

Feb. 10^/i.— One of the feathers was accidentally jiulled out in 
Cock A while the tail was tied up in paper. I do not know how 
this occurred ; I only saw the I'oot of the feather hanging free 
while the length of it was held by the paper. Wlien a gi-owing 
feather is thus pulled out, the horny sheath comes out of the 
deimal socket and i-emains attached to the base of the feather, 

238 MR. J. T. CUNNINGHAM ON [Mar. 17, 

and the end of the sheath is quite soft and exudes a little blood 
and moisture when pressed. This feather was 52-8 cm. or about 
1 ft. 9 in. long. It was one of the anterior tail-coverts, and was 
the only feather ever pulled out of Cock A. 

Feb. Wth. Age 8 months nearly. — On this date, when I was 
stroking the feathers of Cock B, another came out. It w^as one 
of the anterior lateral coverts of the right side, and its length was 
15^ inches (38 cm.). The feather had not ceased growing, but 
showed signs that it was about to cease, as the rhachis at the base 
was stout and thick, and the barbs emerging from the sheath were 
free and downy, as they are at the base of a full-grown feather, 
and never at its more distal part 

Feb. \ith. — Examined the bases of the feathers in both cocks. 
In A the central rectrices had nearly or quite ceased growing, and 
also nearly all the tail-coverts, only two or three still showing the 
basal sheaths. In B two or three of the posterior coverts had 
ceased growing on each side, but a number of the others still 
showed vigorous growth. 

Feb. I5th. — The longest feather in Cock B came out when I 
was stroking the feathers, though I was not pulling at all hard. 
Tlie feather was the central rectrix of the left side, and was in 
growing condition, with bluish pulpy sheath at the base. It came 
clear out of the socket and was not broken off at all. The extreme 
length was 2 ft. 41 in., but about 1 inch of this was in the socket 
before it was pulled out. The total length of the sheath up to 
the point where the feather proper emerged was 3-1- inches (8 cm.). 

Feb. IQth. — Ascertained that in Cock B three tail-coverts on the 
right side had ceased growing and two on the left. The 2nd 
recti'ix on right side had ceased to grow, that on the left nearly so. 

In Cock A the 2nd pair of rectrices had ceased growing, the 
1st or central pair had nearly ceased. Two central coverts, which 
were nearly as long as the central rectrices, had nearly ceased, and 
there were only two others on each side with growth-sheaths. 

There were two rows of coverts in front of the rectrices, anterior 
and posterior, with four or five feathei\s in each i-ow on each side. 
When the feathers are specified individually they are counted from 
the middle line. 

Feb. nth. — One of the posterior lateral coverts on i-ight side in 
Cock B was pulled out today. I was really trying to see if it 
would come out when pulled, as it had completely ceased to grow. 
I found that it required much more force to pull it out than is 
required in the case of a growing feather. The quill was com- 
pletely formed. It was evident that the attachment of the feathers 
was most feeble when growth was ceasing, not when the feather was 
growing vigorously ; while after the quill was completely formed, 
the feather was very firmly held in the socket. The feather was 
19| inches long. 

March 2nd. — In Cock B another feather was unintentionally 
pulled out. It was in full gro^vth, and was one of the lateral 
anterior coverts on the right side. Total length 16^ inches. 


March 4i/i,— Mejisui-ed feathers, exactly four weeks since last 
measurement : — 

Cock A, longest feather 2 ft. 3.J in. 

Cock B „ „ 2 „ U „ 

The longest feathers in A were the two central rectrices, and 
these had almost entirely ceased growing since last measurement. 
They had grown 2| inches in four weeks. 

In B, on the other hand, the longest feather was the central 
rectrix on the right side, wdiich was still growing and was 1 inch 
longer than the rectrices of A. The left rectrix of B, which was 
pulled out, would have been somewhat longer. 

In Cock A there were now only two feathers with growing- 
sheaths, both on the right side, all the rest having ceased to grow. 
in Cock B, on the other hand, there were several coverts with 
growing sheaths, as well as the central rectrix of the right side. 

March 9th. — The two feathers still growing in Cock A were the 
central posterior covert on the right side anil one next to this on 
the same side. They seemed to be about to stop growing. There 
wei-e only five long feathers in Cock A, all the rest being shortei- 
than the coi-i-esponding feathers in B. 

March 10th. — Another featlier came out fi^omCock B today when 
the feathers were stroked ; it was the 1st anterior covert on the left 
side, and, although still growing, showed signs of cessation of growth. 
It was evident that the featliei-s, as noted abo\-e, were easily 
pulled out at this stage. Feather measured 1 ft. 9| in. (55-3 cm.). 

March llth. — Still another feather pulled out of Cock B, 1st 
anterior covert of right side. It measured 1 ft. 8 in. 

March 16th. Age 9 months. — The two feathers which were 
still growing in Cock A seemed now to have both ceased to grow, 
one of them certainly. The only other growing feather in this 
bird was the one on the left side which had sprouted in place of 
the one pulled out. 

In Cock B four of the old feathers weie still growing, namely, 
the right centi-al rectrix, and thi-ee coverts on the left side, besides 
the new feathers sproiiting in place of those pulled out. 

Many of the golden saddle-hackles in B had growing sheaths. 
One came out when they were being stroked the other day, and 
measured 7 inches in length. A few of the feathers of the same 
kind were growing in A, but most have stopped, and these hackles 
altogether were about 2 inches shorter in A than in B. 

Between the saddle-hackles and the long tail-coverts there were 
a number of featheis which may be called transition feathers. 
They were flexible and projected upwards first and then drooped 
in an arch ; the outei- ends were steel-blue, the bases cariied pure 
white down, which was exposed and was very ornamental to the 

Aj^'il 1st. — Measured the feathers, exactly four weeks since last 
measurement : — 

Longest feathers in A, central rectrices, 2 ft. 4j in. 
Longest feather in B, right central rectrix, 2 ft. 8 in. 

Proc. Zool. Soc— 1903, Vol. I. No. XVI. 16 



[Mar. 17, 

The rectrices in A had therefoi'e grown 1 inch in four weeks, 
the rectrix in B 3| inches. The former had ceased to grow some 
time before this date. 

One coveit on the light side in A had still a httle of the blue 
sheath at the base ; it had almost finished growing, and was still 
shorter than tlie four longest central feathers. 

In B the next longest feather was the central covert of the left 

Text-fig. 41 . 

Japanese Loug-tailed Fowl. 
Coct A, photographed April 1903. 

side, which was 1 ft. 1 1 in. long, or 9 inches shorter than the central 
rectrix. The left central rectiix, which had been pulled out on 
Feb. 15th, was growing again rapidly and was now about 4 inches 

In Cock B the left central covert Avas still growing vigorously, 
with long sheath. It had a twist in it and hung in a spiral. Of 
the other two growing coverts on the left side, the outer had 
nearly ceased growing, the other still had a blue sheath. 




1 found that tlie featliers wliicli lind ceased <i;rowiiiy for some 
time were very firmly attached, it was impossible to pull them 
out by moilei'ate force, and I did not wish to pull them out by 
violence. Tliei-e were 8 lonjj^ coverts on each side in both cocks. 

April 7th. —There were siii^ns that more of the feathei's in Cock B 
were coming to an end of their growth. The sheath of the right 
central rectrix was diminishing. The outer of the two n[)per 

Text-fig. 42. 

.lapanese Long-tailed Fowl. 
C;o(,k B, photographed April 1908, 

posterior coverts on the left side had, 1 believe, ceased to grow, 
and the inner, though still growing, was appai-ently aliout to cease. 
The only feather which appeared to be in full growth was the 
central covei-t on the left side. 

April VMh. Aye 10 months. — The right central rectrix in 
Cock B came out today wdien I was pulling the feathers. I had 
pulled it I'ather hard and rathei' frequentlv in the hope of stimu- 


242 MR. J. T. CUNNINGHAM ON [Mar, 17, 

lating its growth, with this result. I am sure it would not have 
grown much more, as its growth was evidently ceasing. 

Total length of the feather 2 ft. 9§ in., of which | in. was 
embedded in the socket. 

April 20th. — On this date the light- coloured hen being bi'oody, 
I put 9 of her own eggs under her to be incubated. 

April 29fA. — Longest feather in Cock B now was the central 
covert of the left side, which was 2 ft. 2 in. long and still gTOwing. 
The covert next to this was also growing. The feathers which 
had been pulled out were all regrowing vigorously, the longest 
being the left central rectrix, which was 7 or 8 inches long. 

In Cock A there was no further growth, except in the anterior 
covert which had been pulled out. 

May 1 1th. — All of the eggs incubated by the light- coloured hen 
were hatched on this date, and all the chicks were vigoi-ovis and 
healthy. My experience of the breed has been in direct opposition 
to the statements in Lewis Wright's account, with regard to their 
vigour and fertility : I have found them exceptionally hardy, 
vigorous, and fertile. 

There was no new variation observable in the chicks, except in 
one which was a rich golden-brown on the head and shoulders, 
instead of light buff colour. All were striped as described in the 
original chicks of the preceding season. 

May 27th. Age 11 months 14 days. — Left central covert in 
Cock B 2 ft. 5 in. long, so that it had only grown 3 inches in four 
weeks. The covert next to this came out on this date when I 
stroked it. It was 2 ft. 2| in. long. It had evidently nearly 
ceased to grow, and was beginning to form downy barbs at the 

June %id. — First signs of moulting noted in the hens ; a good 
many of the breast- and body-feathers on the grovmd from day to 
day, and one primary wing-feather from the light- coloured hen 
found. There was, however, no sign of moulting in the tail- 
feathers of either cocks or hens. Cock A had lately been left 
at liberty with tail free, and none of the feathei"S had been 
broken or lost. 

June Sth. — Hens moulting a good deal, moi-e primaries shed, 
and in some cases the new primary was half developed. The 
primaries in each hen were shed a single pair at a time. 

In the cocks none of the wing-feathers, saddle-hackles, or tail- 
feathers had been shed. 

The chick of this season's brood which was browner than the 
others in the down, showed the same peculiarity in the immature 
plumage, having a yellowish -brown colour instead of the neutral 
brown of the others ; the marking was the same. 

Ju7ie 24:th. Age 1 year 1 1 days. — Left central covert in Cock B 
2 ft. 8 in. long, so that it had grown only 3 inches in four 

July I3th. — The left central covert in Cock B was accidentally 
pulled oiit when I was away fiom home ; the paper in which the 


feathers were tied up liaving come undone, the feather got caught 
in a door and was pulled out. There was some indication that 
the feather was ceasing its growth, as, although the sheath was 
long and soft, the harbs at the base were beginning to be slightly 

The total lengt.h of the feather was 2 ft. IQi in., or 2 ft. 9^ in. 
beyond the socket. 

As this completes the history of the feathers of the first season, 
I will here give a table showing the history of the feathers which 
were pulled out, as it is the further history of these feathers 
which constitutes the most impoi-tant result of my experiment. 

Feathers of fiist adult plumage in Cock B pulled out. 

Mght side of Tail. Ex^^L'^tl. 

Small lateral coveit, growing Dec. 27. 

Anterior lateral covert, nearly ceased Feb. 1 1 . 

Posterior covert, ceased Feb. 1 7. 

Anterior lateral covert, growing Mar. 2. 

Anterior 1 st covert, growing Mar. 1 1 . 

Central lectrix, growing Apr. 1 3 

Left side of Tail. 

Cential rectrix, growing Feb. 15. 

Antei'ior central covei't, growing Mar. 1 0. 

Anteiior 2nd covei't, nearly ceased May 27. 

Posterior central covert, gi'owing J^ily 1 3. 

Of these it is important to note that all but the last two 
sprouted again almost immediately, and continued to gi'ow, 
though not very f;ist, till the general moult. The last two did 
not appear again until the month of Septembei-, that is to say in 
the moulting-season. It is evident, thei'efore, that the reci'escence 
of a feather is influenced by the sea,son of the year. After the 
moult, that is to say in wintei' and spring, there is a general 
tendency to feather-growth, and if a feather is pulled out duiing 
this time a. successor at once begins to grow in its place. After 
the end of April, at any late in the second year of the bii'd's life, 
the activity of feather-growth slackens or ceases, and feathei'S 
pulled out after this time are not replaced till the following 
moulting- period . 

Jtdij 3lst. Age 1 year 1 month 18 days. — Found today that 
Cock A had begun to moult his wing-primaries : there wei'e thi-ee 
or four new feathers growing, and one of them was already half 
its full length. Cock B was in the same condition, but had not 
shed so many feathers. The wing-secondaries were moulting also. 

Ati,g. Ath. — Found one of the outermost shoi-t tail-coverts in 
Cock A growing again, the old feather having been moulted. 
This was the beginning of the moult in the tail-feathers. 

244 MR. J. T. CUNNINGHAM ON [Mar. 17, 

I believe there were 10 tail-coverts on each side in each cock, but 
the two outermost on each side wei-e small and unimpoi'tant. It 
will be seen fi'om the list given above that only eight of the 
covei'ts in Cock B had been pulled ovit, so that eight feathers wei'e 
left, apai't fi'om the I'ectrices, which had completed theii- giowth, 
and could not grow any longei- till they wei-e moulted ; while in 
Cock A all the feathers except one had completed their growth, 
and must necessarily have been moulted befoie they could have 
grown again, 

Aug. llth. — The longest feather in tail of Cock B now was the 
central rectrix of the left side, which 1 measured today and 
found to be 1 ft. 8| in. long. 

The old feather was pulled out on Feb. 15, so that its successor 
had been growing nearly six months, and had grown at the rate 
of little more than 3 inches per month. 

Aug. 16th. — One of the dark-headed hens had moulted all the 
rectrices as well as some of the tail-coverts, so that the i-ectiices 
were shed almost simultaneously. 

Aiog. 24:th. — Found that Cock A had shed another shoi't tail- 
covert, one of the transition-feathers, and also three of the outer 
rectrices : the latter were the 6th and 7th on i-ight side and 6th 
on left, counting from the centre. 

Cock B had shed none of the i'ectrices oi- old tail-coverts yet, 
but this morning when I was stroking the feathers the 1st anterioi- 
covert on the right side came out. The pi-edecessor of this had 
been pulled out on March llth, and it was in full growth. It was 
13f inches in length, and this was the growth of five months. 
There was no reason to regard the loss of this f eathei- as connected 
in any way with the moult, as growing feathei's ai'e not moulted. 

The feathers growing in Cock B in place of those which were 
pulled out in the spring were as follows : — 

Three anterioi- covei'ts on right side. 
One posterior ,, ,, 

One anterior covert on left side. 
Two cential rectrices. 

Sept. 1st. — The outermost rectrix on each side in Cock B came 
out when I was stroking the tail : they were loose and just about 
to moult. This may be regarded as the beginning of the moult 
of the tail, although one of the saddle- hackles and of the inter- 
mediate feathers had been moulted occasionally for some time. 

iSe])t. 2nd. — The 2nd posterior covert on right side in Cock B, 
a full-grown feather, was moulted. It was 1 ft. 7| in, without 
the imbedded quill, 1 ft. 8| in, in exti'eme length, 

iSe2}t. 'S7'd. — 6th rectiix on I'ight side shed in Cock B ; it was 
7 1 inches long without the quill, 

Se2}t. 4:th. — 1st (nearest centre) posterior covert in Cock B 
moulted ; it was 1 ft, 7| in, long. Also one of the outei' anterior 
coverts in Cock A moulted, 

Sept. 5th. — In Cock B three rectrices and one covert on left side 


came out when I stroked the feathers : these were all feathers of 
completed growth, which came out in consequence of the moult. 

In the light-coloured hen, nearly all the rectrices and some of the 
coverts came out when I tried them. The two central rectrices 
in the hen were longer than the others, slightly curved downwards 
at the tip, and speckled with bi'own at the boideis : length 
22'5 cm. (9 inches) without ([uill. The other rectrices were broader 
and black, length 19'5 cm. (8 inches) without quill : the quill was 
1'8 cm. in length. 

Sept. ^th. — More tail-feathei's moulted from Cock B this 
morning. The 2nd anteiior covei't on left side pulled out on 
May 27th ha-d just begun to appear, so that renewed growth was 
defei'red till the moulting-sea.son. 

Sept. 1th. — Nearly all the rectrices and longer tail-coverts which 
had completed their groAvth had now been moulted in Cock B. 
I had pulled many of them out without violence when they were 
loose. By looking at the ba«e of the featheis, I could see when 
they were loose, because the tiansparent hollow quill was pushed 
out and was visible Iteyond the socket. 

In Cock A only four of the rectrices and none of the longer 
covei'ts have been moulted spontaneously. 

The left central rectiix in Cock B, which was still growing, 
measured today 2 ft. Oj in., or had grown not quite 4 inches in 
foui' weeks. 

Sept. \9>th. Age 1 year 2 months. — Today, when taking up 
Cock A to examine him, I accidentally trod upon the gi'owing 
feather the predecessor of which had been pulled ovit on Feb. 10th. 
It was one of the anterior coveits on the left side, possibly the 
first. The feathei- bioke off at the top of the sheath, the latter 
I'emaining in the socket. It was nearly 18 inches long (44"5 cm.), 
and would probably continue to grow from the same base. 

This cock had still six long feathers not moulted, namely : the 
central rectrices, the 2nd pair of rectiices, and two posterior 

In Cock B the fourth anterior covert on the right side, the 
outer of the three growing again, was coming to an end of its 
growth, showing downy barbs at the base, and it would be curious 
to see if it were moulted. 

This fact is worthy of particulai- notice, for it proves that a 
feather which rephices one pulled out before the moidting-season 
does not necessarily continue to grow through that season, and 
it I'aises the question whethei' feathers which are growing when 
the moulting-season begins, necessarily grow longei' or for a 
longer time than those which are moulted in the natural way. 
This question is answered to some extent by the subsequent 

At the end of September I cea«ed to i-eside at Penzance and 
came to London, and having obtained permission through my 
friend Mr. F. E. Beddard, F.R.S., to deposit the fowls at the 
Society's Gardens, 1 took them there on Oct. 1st. They were 

246 MR. J. T, CUNNINGHAM ON [Mar. 17, 

accommodated in two of the pheasants' aviaries. I brought them 
from Cornwall in hampers in a night train, with the tails tied up 
in papers, and none of the feathers were injured. While they 
were at the Gardens I kept the tail of B tied up, while that of A, 
in which there wei-e no long gi'owing-feathers, was left free. Both 
the bii'ds were left at liberty. During the fortnight after their 
ai-rival, all the remaining old feathers in both tails were shed, and 
the moult was thus completed, while in B there were left the seven 
long feathers which had grown in place of those pulled out earlier 
in the year. 

This observation disposes, at any I'ate for these specimens, of the 
supposition that the tail-feathers of the breed are not moulted 
when left to themselves. 

Oct. 11th. — Took away Cock B and the light-colovired hen from 
the Gardens and placed them in a private fowl-house. The left 
central rectrix measured 2 ft. 5g in., showing a growth of about 
5 inches in five weeks. I fixed up a shoi't perch on a stand, and 
at first allowed both birds to fly up to it in the afternoon, then 
later in the evening tied the cock to the pei-ch and unfastened its 
tail till the morning. 

Oct. 22nd. — I intentionally pulled out the fourth of the anterior 
covei"ts on the right side which had stopped growing, in order to 
make the new feather grow again. Thus there were left only six 
long feathers, all growing. 

Oct. 2'dth. — In the morning found the Cock B hanging from the 
perch by the leg which was attached by the piece of tape. 
Generally, if he tried to fly off when he was tied, he was able to get 
back again, but this time the tape slipped down the vertical sup- 
port, and he was unable to do so. In his struggles he had injured 
the longest feather, the left central rectrix, at the base, and it 
broke oW near the top of the sheath. The detached part was 
2 ft. 5-|- in. long. The root of the feathei- Avas uninjured, and the 
basal part continued to grow. 

Nov. 2nd. — Measured the central rectrix of the right side, which 
was now the longest feather: it was 1 ft. 11 in. in length. 

Nov. 15th. — Examined Cock A at the Zoological Gardens. The 
rectrices were nearly full-grown, except the two central pairs 
which were shed last, and which wei'e only a few inches in length. 
The covei'ts wei-e only a few inches in length, and were not likely 
to reach the ground for some weeks. 

Nov. 17th. — This evening when I removed the paper from 
Cock B, I found that another of the long feathers had come out. 
The bird had been free all day, and must have pulled out the feather 
by catching it in some projection when wallowing in the eaith, or 
must have accidentally pulled it out when pi'eening himself. It 
was the second anterior covei't of the right side, and was 1 ft. 9^ in. 
long (53'8 cm.). 

There were now only four long gi-owing-feathers besides the 
left central rectrix which was not lost but broken off. 

Bee. Qth. — I brought away Cock A and the other three hens 


from the Gardens. The new featliers in A hail grown no faster 
than the new feathers in B, in fact they were not quite so long. 
The longest feather in B hnng aliout 2 ft. 8 in. from the perch, 
and all four of thellong feathers were still growing. In Cock A, 
none of the featliers touclied the groimd yet. 

Dec. 9t/i. — Another of the featheis in Cock B pidled out today, 
namely, the central anterioi' covert on the left side. The loss was 
due to accident when the bird was free, not to manipulation. 

Since this date I had kept Cock B tied up nearly every day in 
a sepai'ate house, to prevent any more accidents. The featheis 
that weie pulled out were all sprouting again, and there were three 
long feathers still growing, namely, the right central lectrix, one 
of the posterior coveits on the light side, and the third anterior 
covert on the same side. The fiist two were about the same length, 
that is about 2 ft. 6 in. 

In Cock A, the 3rd pair of rectrices had ceased to grow, and 
were a little longer than the outer lectrices. The 2nd pair of 
rectrices were still growing, and were not much longer than the 
3rd pair. The cential pair weie glowing, and were only al)out 
4 inches long. The tail-coverts were growing, but still very short. 

In Cock B, the 2nd paii- of rectrices were growing, and about 
9 inches long ; the 3rd lectrix on the I'ight side had ceased to 
grow, and the basal sheath was peeling oS", the coi-responding 
feather on the left side was still gi'owing. 

Dec. 21st. — The third anterior covert on right side, which was 
pulled out last spiing and grew again, appeai-ed to have almost 
come to the end of its giowth : the basal barbs were downy, and 
the sheath at the base had almost all peeled off. I had not pulled 
it out. 

Histoiy of Feathers of Second Growth in Cock B, 
to Dec. 28, 1902. 

Bight side of Tail. 

1st Anterior Covert : the preceding feather was accidentally 
pulled out while growing on March 11th, when it was 1 ft. 
8 in. long. It was accidentally pulled out a second time 
while growing on August 24th, when it was 1 ft. 1^ in. long. 
A successor immediately sprouted in its place. 

4th Antei'ior Covert : stopped growing, and formed a complete 
quill a little before October 22nd, on which date it was pur- 
posely pulled out, and its successor spi'outed in its place : it 
was not measured. Originally pulled out December 27th, 
1901, when it was 1 ft. 2 in. long. 

2nd Anterior Coveit : preceding feather pulled out March 2nd 
when 1 ft. 44 in. long: accidentally pulled out again while 
growing on November 17th (1 ft. 9g in. long), and its 
successor sprouted in its place. 

3rd Anterior Covert: predecessor pulled out February 11th, when 

248 MR. J. T. CUNNINGHAM ON [Mar. 17, 

1 ft. 2|- in. long : had almost entirely ceased to grow, but had 
not been pulled out : it was a little over 2 feet long. 

Posteiior Covert pulled out February 17th, 1902, when it was about 

2 feet long. Now still growing, and about 2 ft. 6 in. long. 
Central Rectrix : predecessor pulled out while growing on 

April 13th, when 2 ft. 9 in. long : was still in full growth, and 
about 2 ft. 6 in. long. 

Left side of Tail. 

Central Rectrix : predecessor pulled out while growing on 
February 15th, when 2 ft. 41 in. long: was accidentally 
broken ofl'on October 29th, but continued to grow from the 
base, and the basal portion is now about 7 inches long. The 
total length of the feather, if it had not been broken, would 
have been 3 ft. Oj in. 

1st Anterior Coveit : predecessor pulled out growing on 
March 10th, when 1 ft. 9| in. long: accidentally pulled out 
growing on December 9th, when 2 feet long. Its successor 
immediately sprouted again, and was just showing. 

There were thus in Cock B only three feathers left of the eight 
which had begun to grow before the moulting-season. The two 
longest of these were about 2 ft. 6 in. long, while the third was 
shorter and had ceased growing. 

The other feathei's which began to gi'ow after the moult were on 
the whole longer than the feathers in Cock A ; and the difference 
between the tails in the two birds, the one artificially ti'eated and 
the other left to nature, was sufficiently striking. 

Summary and Conclusions. 

These observations extended only to the 2nd moult, or the first 
shedding of the adult plumage. 

They show that when the feathers are not pulled or artificially 
treated in any way, cai-e being taken to protect them and prevent 
them from being accidentally pulled out, the growth continues 
till about the end of March, when it ceases, the quills of the feathers 
are foimed in the normal way and the featheis are moulted nor- 
mally in the following autumn. 

On the other hand, when the feathers are pulled out in the 
spring, successoi's immediately spi'out in their places, continue to 
grow till the following season, when they go on gi'owing without 
moulting, except in some cases when gi-owth may come to an end 
in the moulting-season. 

In the cock whose feathers were stimulated by pulling, growth 
did not go on at a more rapid rate, but continued for a longer time 
and produced a longer feather. Thus in Cock A, no growth took 
place after April 1st, and the maximum length was 2 ft. 4^ in.; 
while in Cock B, growth continued till July 13th, and the 
maximum length was 2 ft. 9^ in. 


I think it is tolei-ably certain that experiments explain 
the method adopted ))y tlie Japanese in the production of tail- 
feathers ot" extreme length, and that nuich that has hitherto l)een 
mysterious and inexplicable in the matter is now explained. The 
statement that the feathers do not moult or moult oidy once in 
three years, or that the Japanese have a secret method of pre- 
venting the moult, is explained if we assume that the Japanese 
fancier strokes the feather as Mr. iS[)arks stated, and that he either 
delilierately pulls a. feather out when it shows signs of diminishing 
growth, or that the feather is automatically pulled out in the 
process when its growth diminishes ; because, iis I have shown, the 
attachment of the feather is very feeble when the growth is ceasing. 
As a rule, when a feather is pulled out its successor immediately 
spi'outs again, and its growth is aftei'wards continue*! regardless 
of the moulting-season. The results of my experiments in fact 
are in complete agreement with the statement of the matter 
furnished l)y Mr. Sparks. 

The long-tailed cock in its perfection, tlierefore, is neither a sport 
nor a breed, but a, pi'oduct of artificial cultivation ; and the excessive 
growth of the fea.thers is the I'esult of stimulation applied to the 
individual. The most important part of the stimulation is not 
the mere pulling of the feathei', but the extraction of it which 
causes the growth of its successor. 

On the other hand, the method of treatment is applied not to 
any breed at random, but to a particular bleed wdiich includes 
seveial varieties of coloui-, and apparently two vaiieties of comb. 
It can scaicely be supposed that the same treatment applied to 
another breed would produce results at all compaiable, and we 
therefore may conclude that in this special breed there is a special 
and extraordinary tendency to growth in the tail-feathers and 
saddle-hackles. The congenital peculiaiity is evidenced in the 
case of Cock A in my experiment. The question therefore arises, 
whether this congenital peculiaiity ha.s been developed entirely by 
spontaneous variations and selection, or whether it has been 
influenced l)y the excessive giowth artificially induced in every 
generation. It is proba1)le enough that in most cases a cock which 
showed the most rapid and most prolonged giowth was used for 
breeding, but it is l)y no means certain that the cocks so used 
were nevei- subjected to the artificial ti-eatment. The proliabilities 
are rather the other way, that specimens which had been found 
to respond to the treatment were used for breeding. It is a 
significant fact that this is the only In-eed of long-tailed fowls in 
existence, and that the method of treatment applied to it is so 
elaboi'ate and so absolutely artificial, re(|uiring daily attention for 
months and years. If a similar result could have been obtained 
by selection alone, it is difiicult to understand why jioultry fanciers 
in some part of the world have not made the disco\"ery. 

The results of this investigation are remarkably in agreement 
with the theory advocated in my l>ook ' Sexual Dimorphism in the 
Animal Kingdom.' In that book 1 pointed out that wherever 


hypertrophy of organs or parts occurred in one sex only, the parts 
affected were subjected by the habits of the animal, as known by 
observation, to special irritation and stimulation. 

Postscript, added April 25th. — On March 9th, as the left central 
rectrix in Cock B seemed to be ceasing to grow, I tried to pull it 
out, but it broke off at the top of the sheath. The total length 
from the surface of the skin was 4 feet. The right central rectrix 
in the same bird on March 16th was 3 ft. 4 in. long and still 
growing. At present it shows no sign of cessation of growth, 
and has now been growing for more than a year. The appearance 
of the two birds at the present date is shown in the two photo- 
graphs (text-figs. 41, 42, pp. 240, 241). 

2. On some Nudibranchs from East Africa and Zanzibar. 
Part lU By Sir G. Eliot, K.C.M.G., H.M. Commis- 
sioner for the East Africa Protectorate, F.Z.S. 

[Received February 20, 1903.] 

Cbratophyllidia africana, gen. et sp. nov. 

One specimen from near Wasin, E. Africa, in 9 fathoms. 

The living animal was described by Mr. Crossland, who dredged 
it, as of a light greenish-yellow colour on the upper surface, but 
with the foot, branchiee, and under side of mantle, white. The 
back was very hard and smooth, but its most remarkable character- 
istic was the presence of a number of papillae, consisting of round 
or pear-shaped bodies set on stalks. The stalks as well as the 
base and tip of these globes were white, but the middle part was 
black, owing to a dense aggregation of black spots, which, however, 
can be seen to be separate under a lens. The globes were quite 
soft and the stalks flexible ; they shook Avhen the animal was 
moved, but were not observed to execute any spontaneous move- 
ments. The mantle-edge was wavy. 

The alcoholic specimen is of a uniform pale lemon-yellow, the 
black bands of the globes being, however, preserved. The breadth 
across the middle of the back is r9 centim. Unfortunately the 
animal is contracted almost into a circle, but apparently the 
length, when stretched out, must have been about 2*2 centim. 
The consistency of the body is like hard wax, and fragments of 
the mantle, which is ample, could easily be detached with the 
forceps. The whole dorsal surface is a mass of closely packed 
spicules. It bears about a hundred of the stalked globes. They 
are of very varying size ; many ai'e quite minute, but the largest 
is about 3 millim. high including the stalk, and about 2 millim. 
across the ball, which is quite soft and can easily be pressed flat. 
They are distributed over the whole of the back irregularly, and 
not in any pattern, but are perhaps thickest round the mantle- 
edge, including the space in front of the I'hinophores. Both the 

1 For Part I. see P. Z. S. 1902, vol. ii. p. 62. 


iliinophore-pockets aiul tlie anal papilla project ; the edges are 
siiiooth. In the preserv'ed specimen the rhinophores are grey. 

Tlie Inaiichiie are arranged in a circnit interrnpted only hy the 
head and genital p.apilla. They vary in size, hut thougii in places 
long and short branchia' seem to alternate, this cannot he said to 
be the general ride. 

Over the mouth are two tentacles each al)out l*;") millim. long, 
and 1 millim. broad at the base. They are not directt^l side\va}'s 
but straight forwai'd, and being set close together so that the 
division is not visible, they appeal- to form a soi-t of head. They 
aie united at their bases. The mouth is lai-ger than is usual in 
this Older, and though it is suctorial is hardly poiiform. Though 
the animal was dissected only three months after capture, the 
internal organs were already much dried and shrivelled, the spirit 
having appaiently been unable to penetiate the hard integument. 
It was clear, however, that the Imccal organs are of the type of 
Phyllidiopsis rathei- than Phyllidia. The buccal opening led into 
a sausage-shaped tube about G millim. long and 2 millim. broad, 
with muscular walls ti'ansversely stripeil. This passed into a long, 
narrow, coiled tube, which preser\ed the same calibre until it 
dilated into the stomach. Two ample glands (salixary ?) entered 
the larger part of the tube on either side, but were not in any 
way fused with it. The liver was large and undivided behind. 

The central nervous system was enclosed in a thin capsule and 
somewliat concentrated, the cei'ebi-al and pleuial ganglia being 
hardly distinguishable and the pedal ganglia lying beneath them. 
The eyes were large, black, and distinct. The genital mass was 
much hardened, but the two spei-matotheca?, one white and empty 
and the other black and full, weie quite distinct. It was impossible 
to ascertain whether the glans was armed with hooks and whether 
the folds on the dorsal wall of the pericardium (sometimes called 
the pericardial gill) were present, but it is higlily probable that 
the species possesses these family characteristics. 

In virtue of its buccal apparatus this animal belongs to Bergh's 
genus Phyllidiopsis, although the tentacles are not attached 
through their whole length and are I'ather larger than is usual in 
the Phyllidiadje. It is remarkal)le that the genus PhylUdiojisis 
contains one species. Ph. papilligera, which has also black papillae 
on the liack. To me, the presence of these dorsal papillfe seems a 
peculiarity sufficiently marked to merit generic rank. If Echino- 
doris is a genus, why should not Phyllidiad;e which have the same 
peculiarity enjoy the same distinction ? I would propose to call 
the genus Ceratophyllidia, and its characters will be : — Back 
studded, ivith papilla' ; buccal apparatus in the knoirn species 
similar to thai o/" Phyllidiopsis. 

Pleurophyllidiella horatii, gen. et sp. nov. 

One specimen from Wasin, East Africa. Mr. Crossland, who 
captured it, gives the following notes on the living animal : — 
"Three inches long. Mantle edyed with light salmon-colour : it 


ends in two ridges near rhinophores, similarly edged ; front edge 
of velum similarly edged. Back grey, mottled with a darker 
shade, the top of the numerous longitudinal ridges being sprinkled 
with clear black spots. Underside white. Gills also white. 
Rhinophoies longitudinally lamellated, grey in colour, and can be 
contracted or reti'acted, though the pockets do not seem very 
definite or complete." 

The somewhat bent alcoholic specimen measures 3 centim. fi^om 
head to tail, but woidd be at least "5 longer if it were stiaightened 
out. The breadth of the back is 1"5 centim., of the foot "6. The 
foot is long and narrow, pointed behind and truncate in front, 
the corners not projecting. 

No caruncle or nuchal papillae ai-e visible, but it is somewhat 
difficult to reconcile the head-parts of the preserved specimen with 
the desciiption quoted above. According to a rough drawing- 
made from the living animal, it would seem that the mantle-edge 
passes between the ihinophores and forms a sort of velum in front 
of them; but in the preserved specimen it appears to lie behind 
them as in an ordinary Fleurophyllidia, and not to pass through 
them at all. 

The salient character of the genus is that there are no branchiae 
and no trace of a bianchial cleft. Lateial lamella? are, however, 
piesent. They are situated exclusively on the under edge of the 
mantle, and not on the sides of the body. They extend from 
the head to the tail, and are about 30 in number on each side. 
They are irregular in size and shape. Some terminate befoie 
they reach the mantle-edge, and some r-un from the mantle-edge 
only halfway to the body. The genital papilla is 4 millim. and the 
A^ent 1'2 centim. from the anteiior end of the body. Cnidophores 
are distinctly visible round the edge of the mantle. 

The moiTth is ventral, and forms a lai'ge transverse slit, with 
slight indications of a T-shape. The jaws are j^ellow, nairow, and 
united so as to form a shape like a boat. They bear no denticles, 
but thei'e aie a few irregular coarse indentations of the edge, due 
apparently to its being jagged by use. The radula consists of 
about 30 rows, the formula for each of which is abovit 50 + 1 -|- 50. 
The central tooth consists of a squaiish basal plate with a long 
cusp, which bears about 10 denticles on each of its sides. The first 
lateral is laiger than the others and resembles the cential tooth, 
except that the denticles are only on the internal side. The 
second and third laterals ai^e also denticulate ; the rest appear to 
be simply hamate. 

This form appears to be intermediate between Pletvroleura, 
which has neither- branchial clefts nor side lamellae, arrd Pleuro- 
phyllidia, which has both. I have indicated its afiinities by the 
name Pleurophyllidiella. 

B^OLIDIA MAJOR, 11. sp. 

(Bergh, in Sempei-'s Reisen im Arch. Philipp., Malac. Untersuch. 
vol. iii. pp. 778-780, 1880.) 


One specimen from Chuaka, E. coast of Zanzibar, nnder a stone 
between tides. 

The living animal \va.s about 4 centiuietres long. The l)oily and 
appendages were of a uniform greyisli white, with spots of a dull 
opaque white. The whole animal closely resembled a kind of 
detaclial^le sea-anemone which is very common at Chuaka, and 
appears to be sometimes almost free-swimming. 

The alcoholic specimen is 3 centim. long and 1 centim. l)roa(l 
at the widest part, including the cerata. The foot is moderately 
broad, and has fairly long tentacular expansions in fiont ; but 
its most remarkable character is the size and distinctness of the 
anterior gi-oove, which measures 2 millim. across. The upper lip 
is separated into two parts by a deep cleft. Tlie oral tentacles are 
large and very thick. The rhinophores are shorter and studded 
with minute knobs, which, in the preserved specimen at any rate, 
appear not to l)e set in rings. The cerata are much flattened and 
almost leif-like, and the hepatic diverticula within them are 
ramided. They b.^gin at the anterior end of the large peri- 
cardial prominence, and are arranged in about 17 groups on each 
side, each containing about 10 cerata. There are vexy distinct 
gaps between the anteiior groups, and a l)road bare space runs 
down the middle of the back, but towards the end of the boily the 
cerata are huddled together and continue until the extreme tip, 
there being no tail. The outermost ceiuta of all the rows are 
smaller, and the inner considerably larger, but at the base of the 
innermost are frequently quite small, some hardly larger than 
tubercles. The genital oritice is below the first group of cerata, 
and the lateral vent behind the second. 

The specimen was only partially dissected. The jaws are very 
large, colourless and transparent, with a perfectly smooth edge. 
The radula consists of 32 pectinate teeth, very similar to those 
df B. mcebii (see Bergh, /. c. pi. IxxLx. fig. 16), with striations 
under each denticle. They are, however, very much broatler, the 
widest measuring 2 millim,, and the denticles are more irregular 
in shape, being pi-obably worn by use. There are about 150 of 
them on the broader teeth. The three or fom- central denticles 
are generally, but not always, smaller than the others. 

This specimen is clearly referable to Bergh's genus BieoUdia, 
and the difference between it and the type is mainly one of size, 
B. mtddi being only 8 millim. long. The similarity of habitat 
makes one thirdv that this may be merely a full-grown individual 
of the same species ; and we know so little of the variations which 
the radula and arrangement of cerata may present in .'Eolids at 
the different periods of their growth, that I am not prepared to 
reject this hypothesis. Still, the single specimen examined by 
Bergh appears to have been sexually mature, and this being so 
the two animals each present peculiarities amounting to specific 
diflferences: — (1) In B. mcehii the tentacles are said to' be " jibge- 
plattet fingerformig" ; the cerata begin behind the rhinophores and 
are set in rows: in B. major the tentacles are stout and round; 


the cerata begin further back and are set in groups. (2) In 
B. mcebii the reproductive orifice is under the third row of cerata, 
and the vent between the sixth and seventh rows, ahiiost dorsal : 
in B. major the reproductive orifice is under the fii'st group of 
cerata, and the lateral vent after the second. (3) In ^. inajor 
the basal part of the teeth is proportionally narrower than in 
B. 7ncebii, and the denticles are more irregular. 


One specimen captured at Chuaka, May 1902, seemed to be a 
typical Bceolidia major, except for a somewhat more ornate 
coloration. The ground-colour was of a yellowish-white with a 
yellowish-brown pattern, consisting of a series of irregularly shaped 
lozenges containing white spots, down the middle of the back as 
in B. major. The oi'al tentacles were white with green stripes. 
The cerata were white with yellow tips, below which was a bright 
blue band. 


(Bergh, Journ. Mus. Godef. Heft ii. 1873, & Heft vi. 1874; 
Beitr. zur Kenntniss der Aeolidiaden, Theil ii. ; Semper's Reisen 
im Arch. Philipp., Malac. Untersuch. iii. p. 879.) 

One specimen from thereof Jembiani, Zanzibar, 3*5 centim. long 
and 1*3 broad in life. The back was almost entirely covered with 
cerata, so that the body-colour was hardly visible. Most of the 
cerata were very dark green with a blight yellow ling, but the 
innermost were white with bluish tips, with only a ring of dark 
green. The oral tentacles were dark blue, with green bases. The 
rhinophores had four bands of colour, which were, starting from 
the base, greenish brown, white, blue, white. On the head were 
two yellow lines, extending from the oial tentacles to the rhino- 
phores, and the margins of the head and foot were also of a bright 
light yellow. 

The animal was stoutly built. The foot projected considerably 
beyond the body on either side. Its anterior angles were expanded 
into long tentaculai' processes. The head had also two lateral 
expansions, from which projected at right angles the very large 
and conspicuous oral tentacles. In life the rhinophores were 
quite simple and fairly long. In the alcoholic specimen they were 
contracted and somewhat wrinkled. The numerous and thick-set 
cerata wei-e disposed on peduncles. There was a bare triangular 
patch behind the rhinophore, and a narrow bare space down the 
middle of the back, but the cerata folded over the latter so that 
neither it nor the transverse bare areas were visible. There were 
about twenty transverse rows of cerata. The first row at the side 
of the rhinophores consisted of about 1 cerata, much smaller than 
the rest. After the third row Avas a distinct gap, and a smaller 
gap after the fourth. After that the rows were so close together 
that they could not be distinguished superficially. The innermost 
cerata were larger than the others, and sometimes bifid : small 
cerata extended almost to the end of the very short tail. 


The jaws were large, witli a smooth cutting-edge. The radula 
consisted of a single seiies of 13 tiunspai-ent yellowish teeth of the 
shape usual in the genus, viz., pectiniforin, with large irregular 
denticles and small accessory denticles. It was sometimes hard 
to decide whether the I'ather small denticles should he considered 
main or accessory ; Imt tlie average number of main denticles on 
each tooth was 10, and the largest number (in one case only) 13. 
The central nervous system was somewhat concentrated. The 
specimen was only jjartly dissected. 

A new species must, I think, be provisionally created for this 
animal, though the discovery of intermediate foi-ms may perhaps 
render its retention unnecessary. In some ways it is itself a con- 
necting-link between C lonc/icirrha {\nd C. annulata, for the former 
is Siiid to have 7, and the latter 5 denticles on each side of its 
teeth, whereas C. africana has 4, 5, 6, oi- 7 indifferently. It can 
hai'dly be C. annulata, for the difference in colour is too great, 
and besides thei-e is much less bare space on the back. Neither 
can it be C. longicirrha, because (1) the coloiution, though similar, 
is still distinct ; (2) C. longicirrha has the back bare up to the 7th 
row of cei'ata, and some of the cei'ata ai'e veiy long, which is not 
the case here ; (3) the rhinophores are not perfoliate. 

This last point is of some importance for the chai'acterisation 
of the genus. In the present animal the i-hinophoi-es were 
undoubtedly quite simple in life, and in alcohol they ai-e wrinkled, 
though it is still possible to see that they ai-e not really peii'oliate. 
In C. longicirrha, Bei'gh says the perfoliations ai-e 14 or 15 in 
number, and not deep. Of C. annidata he says that the i-hino- 
phores have 12-14 well-marked perfoliations, and that Gari'ett has 
wrongly represented them as simple. But in Semper's ' Reisen,' 
xvii. he states that C. annulata var. affinis has simple i-hinophores, 
and gives as a generic charactei* : " Die Rhinophoi-ien scheinen 
nicht pei'foliirt zu sein." I have not access to part ix. of his 
' Beiti-iige zur Kenntniss der Aeolidiaden,' which j^erhaps explains 
the matter ; but it looks as if the I'hinophores are simple, but have 
a tendency to simulate perfoliations when preserved. 

Pteraeolidia semperi. 

(Bergh, Beitr. zur Kennt. der Aeolidiaden, iii. p. 22, and in 
Semper's Reisen, Malac. Untersuch. vol. i. p. 18 (1870); under 

Four specimens, which seem probably referable to this species, 
were di-edged fi-om 3 fathoms near Chuaka in July 1 901 . The body 
is veiy long, narrow, and vermiform, the lai-gest individual being 
5*5 centim. long and only 3 millim. broad. The ground-colour of 
the body in the living animal is bi'own, with opaque mai-kings of 
veiy light green on the sides and back. The ceiuta are also dark 
brown, with numei'ous thin lines of the same green. The top of 
the head and the ends of the oi-al tentacles ai-e opaque yellowish 
\vhite. The lowei- part of the tentacles bi'own, with three i-ings of 

Proc. Zool. Soc— 1903, Vol. I. No. XVII. 17 


vivid crimson-lake. The body of the rhinophoi'es is brown and 
inconspicuous, but the tips ai-e whitish and have a crimson ring. 
Although the colour of the animal when analysed is as desci-ibed, 
the general effect in most lights is that it is purple with a silvery 

The specimens were active in their movements, and in particular 
the long oral tentacles were very mobile. The anterior margin of 
the foot was produced into two deeply-grooved processes, and its 
sides projected in two laminse along the whole length of the body. 
The cerata, which were not at all caducous, were set on fan-like 
pr-ojections of the dorsal margin, from 13 to 20 in number, on 
each of which were from 10 to 20 cerata. The largest individual 
appeared to have about 640 cerata in all. None of the cei^ata were 
large, but they became more crowded and smaller towards the end 
of the body, where they extended to the tip of the tail. In three 
of the preserved specimens there was a very distinct raised border 
on each side of the back, connecting the fan-like projections, and 
almost resembling the mantle-edge of Dorids ; but in the fourth 
this I'emarkable character was not visible. The oral tentacles wei-e 
ver};' large and long ; the rhinophores, which stand between the 
first clumps of cerata, were small and lamellated, except the tips, 
which were nari-ow and smooth. In the individual dissected the 
radula consisted of a single series of 18 yellowish teeth. The 
separate teeth were much as represented in Bergh's plate (I. c. 
pi. iii. figs. 5, 6, 7), but somewhat more regular in shape, having 
nine denticles very symmetrically arranged on each side of a 
central cusp, but not taking the foi-m of indentations of the cusp 
itself. The cutting-edge of the jaw bore a row of fine but very 
distinct denticles, at the side of which were several I'ows of less 
distinct accessory denticles. No spine or other armature was 
discovered in the I'epioductive system. 

These specimens seem to clearly belong to Bergh's genus Pteraeo- 
lidia. There are some small differences (such as the shape of the 
rhinophores and of the teeth and the lateral ridges) betAveen them 
and his description and plates of Pteraeolidia ( FlabeUina) semperi^ 
but the ridges wei-e absent in one specimen and the other characters 
were slight. It is possible, however, that a comparison of the living 
animals might shoAV a specific difierence. 

Ercolania zanzibarica, n. sp. 

(For the genua Ercolania seeTrinchese, " Un nuovo genere della 
fam. degli Eolididei" Ann. del Miis. di Stor. Nat. di Genova, ii. 
1872; id. " Aeolidsedel Porto di Genova"; Bergh, Beitr. z. Kennt. 
der Aeolidiaden, v. pp. 9-18, Wien, 1878; Yayssiere, Rechei'ches 
sur Moll. Opisthobranches, Marseilles, 1888, pp. 121-128.) 

Two specimens captured at Chuaka, East Coast of Zanzibai-, 
in February 1901. The living animal was rather more than 
2 centimetres long, very fragile and delicate, transparent and of a 
uniform bright gi-een, the hepatic diverticula in the cerata not 


liaving been distinguished by any diflerence or intensity of colour. 
It exactly resenililed a piece of the x-acemose seaweed {Caulerpa) 
on Avhich it was found. 

The length of the alcoholic specimens is 2 centim.; the extreme 
breadth of the l)ack with cerata 8 millim., and of the foot 2-3 millim. 
The rhinophoretj are long and distinctly canaliculate. There are 
no oral tentacles, but two lobes ovei- the mouth. Behind the 
rhinophores are two very distinct black eyes. The cerata are 
clu1>shaped as in Galvina, of varying size, the largest inside. On 
each side of the back are four clumps of about nine cerata each, 
and there is a thick bunch on the tail, which, however, projects a 
considerable distance behind the last cerata. Down the centre of 
the back is a broad bare space, in the anterior portion of which 
is the very large, elongated (not oval) pericardial prominence. In 
front of this and^ fused with it is the vent, a large and conspicuous 
tube. The foot is rounded in fi'ont. 

I dissected one specimen, but was unable to obtain a clear view 
of either the central nervous system or the reproductive organs. 
The latter, as usual in this family, Avere extremely complicated, 
both the prostate and albumen-gland appearing to be extensively 
ramified. The verge was armed with a small spine. The hepatic 
diverticula in the cerata, being colourless, were not easy to dis- 
tinguish, but appear not to be much ramified and to resemble the 
figure of those in Ercolania siottii in Trinchese, pi. ix. fig. 2. 

The mouth-parts, buccal muscles, radula, &c. are of the usual 
ascoglossan type. The teeth are not unlike those oi Ercolania 
viriclh (v. Bergh, I. c. pi. xii. figs. 3 & 4), but the doi'sal surface 
is a simple curve and does not show any depression. The upper 
portion of the radula contains 6 teeth, the lower 27, the last 
members being arranged in a spiral like that depicted in Trinchese's 
plate of B. siottii (pi. x A. fig. 1), from which it may be concluded 
that the individual is young. 

As the specimen presents all the characters of the genus Frco- 
lania, I describe it under that name, but I feel very doubtful if 
the genus is valid. The only characters which difierentiate it 
from Stiliger, viz. that the rhinophores are canaliculate and the 
pericardial prominence elongate and not oval, are surely very 
slight. Yayssi^re [l. c. p. 122) referred to the genus a species 
(fauerea) with entire rhinophores, which is probably in any case 
a Stiliger. 

The animal is not likely to be specifically identical with 
E. viridis Bergh, for the coloration is not really the same, the size 
is much larger, and the shape of the teeth somewhat diflferent. 


258 MR. W. p. PYCRAFT ON THE - [Mar. 17, 

3. Gontribiitions to the Osteology of Birds. 
Part VI. Cuculiformes} By W. P. Pyckaft, F.Z.S., A.L.S. 

- [Received March 17, 1903.] 
(Plate XXII.' & Text-figures 43-48.) 


i. Introductory Remarks, p. 258. I vii. The Pelvic Girdle, p. 280. 

ii. The Skull of the Adult, p. 258. viii. The Pectoral Liinb, p. 283. 

iii. The Skull of the Nestling, p. 268. ix. The Pelvic Limb, p. 284. 
iv. The Vertebral Column, p. 271. x. Summary, p. 286. 

V. The Ribs, p. 274. xi. Key to the Osteology of the 
vi. The Sternum and Shoulder-girdle, Cuculiformes, p. 288. 

p. 275. I xii. Explanation of the Plate, p. 291. 

i. Introductory Remarks. 

Although numerous contiibutions to the osteology of the 
Cuculiformes have been made during the last quarter of a 
century, the present paper claims to be the most comprehensive 
survey of the group from this point of view which has yet been 
made. Nevertheless, much remains to be done before our know- 
ledge is anything like complete on this subject. How much, may 
be gathered from the fact that out of 45 genei'ally recognised 
genera of the Family Cuculidfe, only 20 are represented in the 
osteological collections of the British Museum, and this appears to 
be the lai-gest collection of the group extant. Of the 6 genera of 
the Family Musophagidae, only 2 are represented in our series. 

ii. The Skull of the Adult. 

The skull of the Cuculiformes recalls, on the one hand — through 
the Cuculi — the skulls of certain Coraciiformes, e. g. Coraciidce 
and Buccones ; and on the other — through the Musophagi — the 
skull of the aberrant Gallifoi'm OpistJiocomus. 

Nevertheless the Cuculiform skull may at all times be readily 
distinguished, and by the following chai-acters : — 

The pterygoid is always fi'ee, and aiticulates with the palatine 
— after the fusion of the hemipterygoid — by a more or less 
obliquely transverse joint ; the lachrymal is always present and 
free ; the antorbital plate (prefrontal) is large, and generally (if 
not always) bears an os uncinatum ; the vomer is vestigial or 
wanting ; the palate is indirectly desmognathous ; basipterygoid 
processes are wanting ; and the postorbital processes are small and 
inconspicuous, never extending down to the level of the quadrato- 
jugal bar. 

The skull of the Musophagi may always be readily distinguished 
from that of the Cuculi by the fact that the mesial borders of 

1 For Part V., see P. Z. S. 1902, vol. i. p. 277. 
' For explanation of the Plate, see p. 291. 


'■° s\ T 'as 8 


na 7. 

als fr 



H Gronvold.del Photogravure by Bale iDaniel?, r- 



the palatines ai-e cut away so as to expose the pai'as^ihenoidal 
rostrum. _ 

The Occipital Region. 

The plane of the occipital foramen slopes ol)liquely l)ackwai'ds ; 
the oiifice is either i-eniform or cordiform. In the obliquity of 
this foramen the Cuculifoim skull difters conspicuously from many 
Coraciifoi'in skulls whei'ein the foramen looks directly do^\^l^vards, 
its vei'tical axis lying- parallel with the long axis of the skull. 
The supr a-f or amincd ridge \iihu.ti\\-i\c&\\Q(\, but can be indistinctly 
ti-aced I'unning downwards, on eithei' side, into the processus alse 

Tlie lamhdoidal ridge in the Oucnli is generally well-defined and 
bluntly conical in outline. In some genera, e. g. Crotophaga, 
Centropus^ Gicira, Rhojyodijtes, Rhinococcyx. and Saurothera, its 
extremities bifurcate, sending foi-ward an anterior band aci-oss 
the inferior bordei- of the squamosal to terminate in a pointed 
squamosal prominence ovei-hanging the quadrate ; and a posterior 
bi'anch, which turns downwards and backwards to disapjDear on 
the free edge of the latei'al occipital. "VVhei-e the lambdoidal 
ridge is well-developed, the ci-anial I'oof, \ying immediately in 
front, is more or less flattened, often, indeed, marked by deep 
temporal fosspe. 

In many genei^a the lambdoidal ridge is much less distinct. 
This occurs where the ci'anial roof is sti-ongly arched so as to rise 
above the ridge with a strong curve, and temporal fossae are 
wanting or confined to the sides of the cranium, e. g. Coua^ 
Cacomantis, Chrysococcyx. 

The supi-aoccipital i-egion is marked by strong muscular depres- 
sions, which in the case of Coua, for example, take the form of 
widely-separated oval scars, each surrounded by a swollen border ; 
whilst in others, as in Eudynamys, these depi-essions are only 
divided in the middle line by a narrow ridge, and are bounded 
infei'O-laterally by slightly swollen bullse produced by the lateral 

The latei'al occipital wing is j^roduced downwards on either side 
into a small or medium-sized jyrocessus alee exoccipitalis inferior 
(paroccipital process). 

In the Musophagi the lambdoidal ridge is fairly well-defined in 
Schizorhis, but only vei'v imperfectly so in Turacus. 

The i-idge in Schizorhis is formed by the scooping-out, so to 
speak, of the supi-aoccipital to aftbrd depressions for the insertion 
of the cervical muscles. On either side, and inferior to these 
depressions, are two rather conspicuoiis bulla? lodging the posterior 
and horizontal semicii-culai- canals of the ear. The distal ends of 
the lambdoidal ridge tei-minate before reaching the fi-ee edge of 
the lateral occipital wing. The 2»'0cessus alee exoccipitcdis ivfeHor 
is only feebly developed. 

The lateral or exoccipital wings are not gi'eatly developed in 
the Cuculiformes. 

260 MR. W. p. PYCRAFT ON THE [Mar. 17, 

The Cranicd Roof. — The cranial roof is never marked by supra- 
orbital grooves. 

The parietal region, in certain genera of Cuculi, is marked by- 
deep temporal depressions (temporal "fossae "), which may meet in 
the middle line so as to form a faint sagittal ci-est, e. g. Rhojjodytes, 
Piaya, Rhinococcyx, and Rhcmiphococcyx ; or they may be divided 
by a low broad ridge, as in Geococcyx, Taccocoiuc, Satf^rothera, 
Exidynamys, Guira, Centropus, and Crotophaga. In Goua, 
CoGcystes, Guculus, and Gacotnantis the temporal fossa is either 
barely perceptible or confined entirely to the lateral aspect of the 
cranium. Geococcyx, it should be remarked, presents an inter- 
mediate stage in this character ; the width of the ridge dividing 
the right and left fossae being extreme, so much so that they 
barely pass beyond the latei'al aspect of the skull. 

The interorbital region is moderately wide in all the Cuculi, 
save in Saurothera, but even in this genus it is not conspicuously 
narrow. In the middle line the region may be marked with a 
distinct furrow as in Goua, or even by a low ridge as in Eudynamys. 
The interorbital region is bounded anteriorl}- by the lachrymals ; 
these do not develop outstanding supraorbital processes, and are 
not conspicuous from this aspect of the skull. There is no trace 
of the fusion of the fi'ontal and rtasal bones. 

In the Musophagi the temporal fossae are not strongly marked, 
and are confined to the lateral aspect of the skull. The parietal 
region is more rounded than in the Cuculi ; and in Tttracus is 
marked by a median furrow, so that the foi'm of the cerebral 
hemispheres is distinctly indicated. In other I'espects this region 
of the skull resembles that of the Cuculi. 

The Base of the Skull. 

The basitemporal plate, in the Cuculi, is flattened and triangular 
in shape, and with the apex lying immediately beneath the 
Eustachian apertures. Posterioily it is bounded by a pi'econdylar 
fossa, which, though generally barely perceptible, in Eudynamys, 
Gentrop'iis, and Geococcyx is fairly deep. The lateral angles of the 
base are truncated, and, turning sharply downwards, ci'oss the 
paroccipital notch to form the lower portion of the rim of the 
mouth of the tympanic cavity. In some genera, e. g. Goua, Gidra, 
these downwardly-directed angles are very prominent and simulate 
mammillary processes. 

The two sides of this tiiangular plate have fused with the 
ossified connective tissue foiining the anterior wall of the recessus 
tympanicus anterior. Consequently the Eustachian grooves have 
been converted into tubes opening directly above the apex of the 

The parasphenoidal rostrum bears slight vestiges of basi- 
pterygoid processes in Eudynamys only among the Cuculi. 

The rostrum is relatively long, and curves slightly upwards. 

In the Musophagi the basitemporal plate, like that of the 


Cuculi, is triangular ; l)iit it tlifters from the Cuculine plate in that 
it is somewhat swollen, a feature which is especially well-marked 
in Schizorhis. The two sides of the triangle are produced hack- 
w^ards over the paroccipital notch to join the lateral occipital wing 
of the tyinpanic cavity : thus they come to project beyond the 
base of the triangle, though they are less conspicuous tliau in 
some of the Cuculi. There is a barely perceptible precondylar 

The Lateral Aspect of the Cranium. 

The tympanic cavity is oval or oblong in form and of moderate 
size. It is bounded in front by the quadrate, behind by the 
lateral occipital tympanic wing, below by the basitemporal jilate, 
and above by the squamosal prominence and by the head of the 

Within the mouth of this cavity lie several smaller apertures. 
The largest of these is that of the recessus tympanims anterior. 
Immediately below this is the mouth of the Eustachian tube. 
The foramen ovale and foi'amen rotundmn, three foramina of the 
sinus petrosus, and the mouth of the posterior tympanic recess 
open into the cavity by a common aperture — tliefenestral recess — 
which is very small, and lies mesiad of the articular surface for 
the otic head of the quadrate. The mouth of the posterior 
tympanic i-ecess is completely concealed, and can only be made 
out by cutting away its outer wall and passing a bristle through 
from its cavity into the common apertui'e of the foi'amina in 
question. In some birds, e. g. Falco, the posterior tympanic recess 
communicates with the tympanic cavity by two apertures— one 
caudad of the foramina ovale and rotundum, and lying within the 
fenestra through which these are appi'oached ; and the other external 
to this fenestra, separated therefrom by a bony column, and lying 
immediately beneath the articulai- surface for the otic head of the 
quadrate. This external aperture is wanting in the Cuculiformes. 
Immediately above the head of the quadrate, and between the otic 
and squamosal heads, will be found the aperture of the superior 
tympanic recess, which is reduced in this group to extremely sinall 
dimensions. The foramen of the 7th (facial nerve) opens also into 
the tympanic cavity between the otic articular surface for the 
quadrate and the lim of the anterioi- tympanic recess. 

The squamosal and otic articulai- surfaces for the quadrate form 
a dumbbell-shaped area immediately in front of the aperture of 
the superior t}-mpanic cavity. The actual articidar suiiaces are 
formed by the expanded ends of the dumbbell only. Where the 
pneumatic area is ver}^ large, as in the Striges and Falconiformes 
for example, it breaks thi-ough the connecting poi'tion of these two 
articulai- sui-faces so that they liecome quite isolated. 

The Squamosal Prominence.— The size of this prominence 
depends largely ujton the development of the temporal fossa. 
When this is deep the prominence is large ; but when shallow the 
prominence is quite inconspicuous. Its anterior angle is produced 

262 MR. w. p. PYCRAFT ON THE [Mar. 17, 

forwards into a more or less well-developed spine, the p7-ocessus 
zygomaticus squcmnosi. The base of this angle serves for the 
articulation of the squamosal head of the quadrate. This zygo- 
matic process is especially well-developed in Cuculus, Crotophaga, 
and Geococcyx. It is small in Gidra and Goua, for example, 
and in the Musophagi. The j^'^^ocessus articularis squamosi is 

The temporal fossse are never very deep, but they may extend 
inwards to within a short distance of the middle line. The 
variations in the extent of these fosste have already been dealt 
with {supra, p. 260). 

The trigeminal foramen in the Cuculi lies immediately above 
the mouth of the anterior tympanic recess, on a level with, and in 
front of, the otic ai-ticular surface for the quadrate. In Turacus 
this foramen is divided by a median vertical bar of bone. 

The orbito-nasal foi-amen (v^) appears to be definite only in 
Geococcyx. In other Cucidinoi and Ihosophagioue the alisphenoidal 
wall only partially ossifies, and consequently this and other 
foramina aiound the optic foramen disappear in the dried skull. 

The orbits are large, and roofed by moderately broad over- 
hanging ledges formed by the frontals, and to a slight extent by 
the lachrymals. The orbito-s2)henoid does not ossify. 

The interorhital septttm in the Cuculi is genei-ally pierced by a 
large fenestra, which attains its maximum in Dromococcyx and 
Centropus. In Grotophaga, Goua, Piaya, Rhamphococcyx, Rhino- 
coccyx, Rliopoclytes, and Zanclostomus, however, the fenestra is 
greatly reduced. 

In the Musophagi the fenestra is large in Turacits, small in 
Schizmdiis. I have not had an opportunity of examining other 

The oi-bit is bounded in front by the antorbital plate (p. 263). 

The lachrymal in the Cuculi is moderately large, with subequal 
supraorbital and descending processes. The form of this bone 
and its relation to the frontal and nasal bones vary considerably 
however. In Rhamphococcyx calorhynclms only, apparently, is it 
attached merely to the nasals. The orbital process looks entirely 
forwards, instead of upwards : that is to say, it forms a shield 
for the fiont of the eye instead of an ovei-hanging ledge therefor ; 
the descending pi-ocess is slender. The frontals rest on the inner 
angles of the orbital process of the lachrymal on either side, and 
appear, in the adult skull, to be cut ofi' from the nasals and pre- 
maxillary process by a shai'ply-defined groove, which runs across 
the skull fi'om the hinder edge of the orbital process up the 
lachrymal on the one side to that of the other, and simulates a 
nasal hinge. Rhinococcyx curvirostris appai'ently closely resembles 
Rha'mphococcyx, but the lachrymal being completely fused with 
the nasals and the frontal behind it, the precise relations of the 
various regions cannot be made out. 

In othei- Cuckoos, though exhibiting a distinct tendency to 
form an anteiior shield rather than a jDent-house ledge, the 


lachrymal extends backwards for some distance over the fiontals. 
In Eadj/namys the fi-ontals ai-e deeply notched to receive the 
orbital process of the lachiymal, which is more or less diamond- 
shaped. In other genera the orbital process of the lachrvnial 
may be semilunai' or rod-shaped. Whei'e the antorliital process 
is very lai-ge, the descending pi'ocess of the lachiymal is always 
more or less markedly degenei'ate. In Cuculus it has disappeai-ed 
altogether, wliilst the orbitnl pi'ocess has fused with the frontal 
and nasal bones. In Zanclostomus javanicus it is I'ednced to a, 
mere vestige. The descending process is unusually large in 
Crotopluiga, one of the genei'a, it will be i-emembered, with a small 
antorbital pi'Ocess. 

Geococcyx has the lai'gest lachiymal of all the Cuckoos. It is 
nearly columnar in form, flattened antero-posteiiorly, and deeply 
gi'ooved immediately above the quadrato-jugal bai'. 

The lachrymal in the Musopliagi in its genei'al sha])e closely 
resembles that of the Cuculi : the oi'bital process is, however, 
inclined more upwards. The descending process, judging from 
what obtains in Schizorhis and Jlnsophac/a , is faiily strongly 
developed. In Schizorhis the fi-ee end of the descending process 
is twisted in itself, i-ests upon the quadrato-jugal bar, and ai-ticu- 
lates by its postero-inteiiial angle with the enormous ossicidmn 

The lachrymo- nasal fossa varies considerably in size. Among 
the Cuculi, it may be described as large in Eudynamys, Centropus, 
Gidra, and Crotophaga. In Coua, Rhamj^hococcyx, and Geo- 
coccyx, for example, it is extremely reduced by the encroachment 
of the lachiymal. 

The Ethmoidal Region. — The onesethmoid, both in the Cuculi 
and the ]\Iusophagi, terminates abrupth' immediately in front of 
the antorbital plate. The horizontal plate formed by the expan- 
sion of its dorsal bordei- is but feebly developed. 

The antorbital plate (prefrontal), which, with the lachiymal, 
bounds the orbit in front, is, in some Cuculi, e. g. Cacontantis, 
Centrojms, Coua, C'occystes, Cacidus, Eitdynamys, Geococcyx, 
Fiaya, I'accocoua, and Saurothera, conspicuously large, quadrate 
in form, and has its dorsal liordei- piei'ced by two foi'amina — an 
inner for the passage of the olfactory nerve, and an outei' for the 
orbito-nasal (v^). 

In other foims, e. g. Gaira, Crotop)haga, the antorbital plate, 
though large, has the external lateral and inferior borders 
deeply excised. In such cases the orbito-nasal foramen described 
above is repi-esented by a notch. Attached to the inferior border 
of this plate is a vestigial ossiculum palatinum, which, however, is 
generally wanting in dried skulls. 

In the Musophagi the antorbital plate is greatly reduced, 
forming but a triangular process projecting from the mesethinoid. 
From its inferior border depends a well-developed ossicxdum 
lachrymo -palatinum. In Schizorhis tiiis bone is relatively 
enormous, projecting downwards to aiticulate by a strap-shaped 

264 MR. w. p. pycRAFT ON THE [Mar. 17, 

limb with the outer border of the palatine. Its further relations 
with the lachrymal will be discussed presently. 

The nasal septum is completely ossified, and the external nares 
are in consequence imperforate. The ectethmoidal region pi-esents 
several points for comment. By the ossification of the alinasal 
wall, the form of the external nares is greatly changed in many of 
the Cuculi. In consequence of this ossification, the space between 
the premaxillary and descending processes of the nasal becomes 
obliterated. Thus in Geococcyx the nostrils take the form of 
an oval aperture lying just cephalad of the middle of the beak. 
In RhampliocoGcyx this aperture is still further reduced, and 
consists only of an elongated slit lying at the base of the beak 
above the tomium. In Rhinococcyx the aperture is nearly circular 
at the base of the beak, and bounded by a deep groove in front. 
The alinasal ossification is, however, impei'fect, inasmuch as a 
small round hole is left in the angle betAveen the premaxillary 
and descending processes of the nasals. In Taccocoua this 
superior foramen and the aperture of the nostril have blended. 

In Coua the floor of the anterior ]'egion of the nasal fossa is 
raised up into a sharp longitudinal I'idge. 

In the Musophagi the foiiii of the external nostril diifers con- 
spicuously from that of the Cuculi. It is situated slightly distad, 
or proximad, of a line passing through the middle of the beak, and 
is renifoi-m in shape. In Titracus it lies distad, in Schizorhis 
proximad of the middle line. In both genera, just within the 
mouth of the aperture, lies a well- developed concha vestibidum. 
This is especially large in Schizorhis. Furthermore, the genus is 
peculiar in that immediately distad of the concha the nasal septum 
is pierced by a minute foramen. 

The Cranial Cavity. — The metencej)halic fossa is somewhat 
shallower in the Musophagi than in the Cuculi. The internal 
auditory meatus in the Cuculi is sharply defined, strongly con- 
trasting in this respect with the Musophagi, in which it is but 
ill-defined. The vagus foramen, which lies caudad and ventrad of 
the meatus, is small. The orbito-nasal forameoi (v^) pierces the 
outer superior rim of the fossa near the outei- angle of the dorsum 

The cerebellar fossa is relatively smaller in the Cuculi than in 
the Musophagi, and in both it is relatively smaller than in many 
other groups, e. g. Falconiformes. In the Cuculi the floccular 
fossa is very shallow, a.nd lodges a deep slit-like vertical 
depression. In the Musophagi this fossa is reniform, deeper, and 
lodges a deep and wide depression which gives the reniform 
character to the whole. 

The mesencephalic fossa is fairly sharply defined in both groups. 
Among the Cuculi, this is especially noticeable in Geococcyx. Its 
floor, in both groups now under consideration, is pierced by the 
trigeminal foiumen. The diflference in position of this foramen is 
worth noticing. In the Falconiformes it lies, together with the 
orbito-nasal (v'), in a pit carved out of the superior border of 


the metenceplinlic fossa, midway between the sella turcica and the 
floccular fossa, and is overhung by a sharp ledge formed by the 
inferior border of the metence'phalic fossa. 

The pitidtarj/ fossa is large, deep, and tubular. The hinder 
border of the fossa— the dorsum seZ^re— forms a tumid lip. 
Anteriorly the fossa is bounded by the 2)re-piti(.itan/ ridge, which 
forms a broad triangular optic platform. This platform is carried 
far forward into the interorbital septum, rising gently in its 
course, and then turning abi'uptly backwards to terminate at the 
p7-e-optic ridge, which may be traced into the tentorial ridge. 
The form of the optic platform contrasts strongly with that in 
some otlier types, e. g. Accipitres. The tentorial ridge in the 
region of the pre-optic platform is but feebly developed. On the 
parietal wall, however, it becomes tolerably distinct, especially in 
Geococc]/x and the Musophagi. In these, on its Avay to the median 
falx it meets, near the crest of the epiotic, the sharply-defined 
internal border of the mesencephalic fossjx, and forms therewith 
a prominent angular boss of bone, especially well marked in 
Geococcyx. The'bony falx is not very strongly developed. 

The ' oculo-motor (iii.) and abducens (vi.) nerves leave by a 
common aperture, foi'ming a deep gi'oove across the posterior 
angles of the optic platform. 

The cerebral fossai lie quite in front of the cerebellar fossa, and 
are more flattened dorso-ventrally in the Cuculi than in the 

The olfactory fossfe are relatively feebly developed. 

The Fremaxilla. 

The external, as distinguished from the palatal, portion of the 
upper jaw is made up mainly by the premaxilla. This region of 
the jaw, in the Cuculi and Musophagi, varies considerably in form. 
iSTormally, it may be described as about as long as the cranium. 
In Croiophaga it is slightly longer than this, in Geococcyx and 
Sanrothera very much so. Tj^^ically, it may be described as 
hemici'escantic ni outline, depressed from above downwai-ds, and 
tapering from the base forwards. In Geococcyx and Sanrothera 
it is long and pointed; in Croiophaga surmounted by a sharp, 
high keel ; in the Musophagi more or less inflated. 

The premaxillary poi-tion of the jaw is slightly decurved at the 
tip, and the palatal surface is level with the culmen. The nasal 
processes of the premaxilla fuse completely with the nasals. 

A nasal hinge occurs in Turacas among the jMusophagi, but is 
wanting in the Cuculi. 

The Maxillo-jugal Arch. 

The maxilla is indistinguishably fused with the premaxilla. 
The maxillo-palatine processes are expanded horizontally, and 
meeting the ventral border of the nasal septum in the middle line, 
foi'm an indirectly desmognathous palate. 

266 MR. w. p. PYCRAFT ON THE [Mar. 17, 

Among the Cuculi, the least sjDecialisecl palate is probably to be 
found in Coua and Etidynamys. In Goua the maxillo-palatine pro- 
cesses are widely separated and spongy in character ; the consequent 
palatal vacuity is filled by the nasal septum, which is also spongy 
in character, and, fusing with the maxillo-palatine on either side, 
forms an indirectly desmognathous palate. Eudynamys closely 
resembles Coua in this respect, but the bony tissue is less spongy, 
and the fusion between the palatine processes and nasal septum is 
more complete. Cioculus, Gfibira, and Gentropus resemble Etidy- 
namys. Further specialisation of the palate is seen in the still 
further obliteration of the boundaries between the maxillo-palatines 
and the nasal septum, and the tendency to shorten and broaden 
the palatines. Forms like Geococcyx and Rhinococcyx show how 
these modifications have come about, whilst in Rhamfhococcyx we 
may see the maximum develoi^ment of these peculiarities. 

In the Musophagi the maxillo-palatines never appear to com- 
pletely coalesce in the middle line, nor does the nasal septum 
descend to the level of the ventral surface. The palate is never- 
theless desmognathous, since this septum fuses with the dorsal 
surfaces of the maxillo-j^alatines. 

No separate elements can be distinguished in the quadrato-jugal 

In the Cuculi the quadrato-jugal bar is almost or quite con- 
tinuous with the tomium of the maxilla : in the Musophagi, on 
the contrary, it I'ises considerably above the level at its distal end. 
This difierence is due to the fact that in the Cuculi the floor of 
the maxillo-j)alatine process lies low, being only just raised above 
the palatines at its postero-external angle, whilst in the Muso- 
phagi this region is raised high above the palatines. 

The Vomer, Pcdatines, and Pterygoids. 

The vomer is absent in the Cuculi, vestigial in the Muso- 
phagi. In the latter group it is spicular in form and fused with 
the palatines, which it joins by means of a pair of very short 
limbs. Anteriorly it touches the nasal septum. 

The palatines, in the Cuculi, vary in shape. In the less 
specialised forms, such as Eudynamys, Goua, Guira, they are 
moderately long, and exposed slightly outwards. Anteriorly, from 
the level of the forward face of the antorbital plate to the point of 
fusion with the maxillo-palatines, the shaft of each is rod-like ; 
cephalad of this point the bone spreads out into a moderately 
broad blade, the postero-external angles of which are rounded off, 
whilst the mesial border is produced ventrad to form a more or 
less pronounced keel. In Guculus there is but little difference 
in the width between the anterior and posterior moieties, and the 
shaft is nearly straight. 

In the heavy-billed forms like Rhamiphococcyx and Rhinococcyx 
the palatines are relatively shorter, and have the maxillo-palatine 
extremity laminate instead of rod-like. The skull of Scythrops, I 
regret to say, is not contained in the National Collection. 


In all the Cuculi, the palatines meet one another in the middle 
line immediately heneath the pavasphenoidal imtrum, whieh they 
gi-asp laterally through their fusion with the hemii)tery<i()ids. 

In the Musophagi the palatines are relatively longer than in 
the Cuculi, from which they also differ in that they trcper, instead 
of broaden, rapidly as they approach the pterygoids. Furthermore, 
they do not meet below the parasphenoidal rostrum, but, on the 
contrary, are separated one from another by the whole width of this 
rostrum '(PI. XXII. fig. 2). The dorsal surface of the pterygoid 
end sends up a long and deep, incurved keel, the antero-internal 
angles of which fuse with the vestigial vomer. 

The ■pterygoids, in the Cuculi, are moderately long and straight, 
and in some, e. g. Crotop/taga, Rhhiococcyx, G'nira, Taccocoua, 
have the dorsal border raised up into a high, sharp crest, the 
palatine end of Avhich embi'aces the parasphenoidal rostrum, whilst 
in Eudynamys, Coiuc, Geococcyx, and Fiaya, for example, this 
crest is absent. The palato- pterygoid articulation is in the form 
of an obliquely transverse hinge-joint, permitting only a lateral 
motion. Basipterygoid facets are absent. 

The hemipterygoid element of the pterygoid is conspicuous only 
in Taccocoua and Geococcyx. Herein it forms a continuation of 
the dorsal crest of the shaft of the pterygoid, and rests on tlie 
palatine at a slight angle. Secondary fusion between the hemi- 
pterygoid and the main shaft of the pterygoid Avould reproduce 
exactly the conditions of the palato-pterygoid articulation which 
obtain in the Bucconidce and 2[omoiida'. 

In the Musophagi the pterygoids are relatively short, somewhat 
twisted, rods, bearing vestiges of basipterygoid facets. They 
articulate with the palatines as in the Cuculi. 

The Mandibles. 

The mandible in the Cuculi has a short, blunt angular process, 
and a modei-ately long internal angular process. 

In Eudynamys, Citcidus, and Gtoira the lumi ai'e pierced by a 
long lateral vacuity, which is partly closed by a long and slender 
coronoid. In Coica the coronoid tei'minates midway aci'oss this 
vacuity ; Avhilst in Taccocoua iind Centropus this vacuity is quite 
open, the coronoid forming its venti-al bordei'. In Fiaya, A'hino- 
coccyx, and Rhampliococcyx the ramal vacuity is absent. In 
Geococcyx it is partly closed anteriorly by the hinder end of the 

The mandible of Cucidas possesses one conspicuous feature in 
the presence of a triangular flange of bone formed by the deflection 
of the superior border of the I'amus at the point corresponding, 
in the living bird, to the gape, in the skeleton to the region 
imniediately behind the lachrymo-nasal fossa. The flange, 
especially conspicuous in Cucidus canorus, is also fairly distinct in 
Cacomantis, and is tracesible in Fiaya and Fhanvjihococcyx. The 
internal angulai- piocess is well marked in all the Cuckoos, but is 
especially so in Coua and Centropus. 

268 MR. ^Y. P, PYCRAFT OjS^ THE [Mar. 17, 

In the Musophagi the angulay' is sharply truncated posteriorly, 
and the internal angular process is short and blunt. 

In Schizorhis the lumal vacuity is closed posteriorly, in Turacus 
it is closed completely, by the coronoid. 

The Hyoid. 

Unfortunately the hyoid bones have been preserved only in one 
or two of the skeletons of this group in the Museum Collection. 
Years ago, when these skeletons were made, according to the 
prevailing custom, the hyoid bones were not regarded as of value. 

In Scythrops the basihyal is slender and rod-shaped ; the 1st 
' basibranchial triangular, with concave sides, the 2nd of medium 
length, styliform and tapering. The ceratobranchial and epi- 
branchial are subequal in length and offer no points for special 

In the Musophagi the basihyal bones are reduced to mei-e 
vestiges. The 1st and 2nd basibi'anchials are short and fused. 
The ceratobranchials are only slightly longer than the epi- 

iii. The Skull of the Nestling. 

The ISTational Collection of nestling skulls of this group is 
extremely limited, containing only one skull of Cuculus canorus 
and one of Geococcyx calif ornianus. A series of skulls of nestlings 
and of immature individuals of various species of Cuckoos, as 
well as of Plantain-eaters, would be a useful addition to the 
Collection. At present the latter contains no skeletons of nestling 
Plantain- eaters and few skeletons of adults. 

a. Cartilage-hones. 

The hasioccipital is somewhat linguiform in shape, and widens 
gradually from behind forwards. It is bounded on either side by 
the latei'al occipitals, in front by the basitemporal plate. Its 
posterior border forms the greater portion of the occipital condyle. 

The exoccipital, or lateral occipital, viewed externally, takes the 
form of a broad horizontal plate bounding the hasioccipital on 
either side, and extending backwards and outwards, expands into 
a fan-shaped plate, the superior half of which is wedged in 
between the supraoccipital, parietal, and squamosal, whilst the 
ventral, or downwardly directed, moiety forms the posterior 
boundary of the tjanpanic cavity. The lateral segment of the 
foramen magnum is formed by the horizontal plate of the lateral 
occipital, which also, by the way, forms the extreme outer angle 
of the occipital condyle. The fact that the exoccipital comes into 
contact with the parietal is noteworthy : a similar relationship 
obtains also in Dromceus among the Palseognathas, and will 
doubtless be found among seveiul other ISTeognathine forms. The 
exoccipital is only just visible, from the cranial cavity being 
ovei'laid by the opisthotic. 


The supraocclpital is extremely short untero-posteriorly, and 
is deeply cleft in the miihlle of its superior border. Its external 
lateral border fuses with the lateral occipital ; Avithin the cranial 
cavity it is bounded by the epi- and opisthotic. The small size 
of the supraoccipital recalls the skull of the Owls, but this of 
course is but a coincidence. The character will doubtless be 
found to obtain in the Kingfishers, Bucconidje, and Capitonidfe, 
which have many characters in common Avith the Cuckoos. 

The prootic does not appear externally. Within the cranium 
it occupies considerable space, forming the floor of the mesen- 
cephalic fossa, as well as a considerable portion of the lateral walls 
of the basin-like metencephalic fossa. It entirely excludes the 
squamosal from the cranial ca\-ity. 

The epiotic is only partially ossified, and in such a way that its 
boundaries cannot be made out. 

The opisthotic has fused completely with the prootic. 

The basisphenoid is not visible externally, being underlaid l)y 
the basitemporal plate. Concerning its internal boundaries, 
nothing satisfactory can be gathered from the two skulls in the 
National Collection, the younger being damaged, Avhilst in the 
more adult skidl it has fused with the neighbouring bones. 

The cdisphenoid, in the younger of the two skulls, is not yet 
completely ossified. As a result, between the external ventral 
angle and the squamosal there is a wide gap, _ Avhich _ extends 
inwards below the inferior alisphenoid border, dividing its outer 
moiety from the basisphenoid. This gap, when viewed from 
without, is seen to be filled up by the prootic. Its supero- 
external angle is produced outwards to form the postorbital 

The orbito-sphowid is not yet ossified in these skulls. 

The p)resphenoid has fused Avith the basisphenoid. 

The meseihmoid, in the tAvo skulls noAV under consideration, 
is yet incompletely ossified, forming but a linguiform plate ; the 
interorbital septum formed by the backAvard extension of the mes- 
ethmoid having been represented only by cartilage. The anterior 
border of the Hnguiform plate is shai-ply truncated so as not to 
extend beyond the level of the free end of the parasphenoidal 
I'ostrum beloAv and the anterior extremities of the frontals 
above. This truncation occurs at the cranio-facial fissure, Avhich 
has cut the mesethmoid into tAvo parts : the one forming the 
linguiform plate just described, Avhich ultimately forms the 
interorbital septum ; the other, the septvm nasi, Avhich in these 
skulls is yet cartilaginous. 

The cranio-facial fissure appears to be peculiar to the neo- 
gnathine (Carinate) skull ; but traces thereof are apparently to be 
met Avith in the Paheognatha? (Ratita?), inasmuch as, in the skulls 
of nestlings of Dromceus and Rhea in the Museum Collection, the 
ossification of the mesethmoid commences, as in the Neognatlue, 
by the formation of a more or less linguiform plate, and this has 
its superior border deeply excised, at a point exactly corresponding 

270 MR. W. p. PYCBAPT ON THE [Mar. 17, 

to the vei-tical face of the posterior division of the neognathine 
mesethmoid, that is, exactly in fi'ont of the antoi-bital plate of the 
adult. To make the fissui'e complete, the incision would have to be 
continued downwai-ds so as to completely bisect the plate, whilst 
the rostrum would have to terminate at the free edge of the 
posterior segment of the plate. In the adult skulls the incipient 
fissui'e remains, placing the right and left olfactory chambei'S, in 
the di-ied skull, in communication. The foi-mation of a complete 
cranio-facial fissure is correlated with, and pei-haps consequent on, 
the reduction in the length of the parasphenoidal rosti'um, which 
in all the Palteognathse is of great length. In a skull of a half- 
grown Casuarms scdvadorii it is, however, relatively much shorter 
than in an adult C. aitstrcdis, and much overhung by the mes- 
ethmoid. Whether this peculiarity obtains also in the adult of 
this species I am unable to say. 

The olfactory cavities do not extend backwards in the Cuculi 
or Musophagi, so as to lie on either side of the mesethmoid, as in 
the Tubinares for example ; the antorbital plate ai-ising in the plane 
of the cranio-facial fissure. 

The quadrate does not differ materially from that of the adult. 

The articidare is still distinct. 

b. The Membrane-bones, 

The 2X(,rietal is oblong in shape, its anteiior and posteiior 
borders sinuously curved, its mesial boi'der sti'aight, and its 
extei'nal latei'al border being slightly convex. It is bounded in 
front by the parietal, behind by the supraoccipital, and laterally 
by the sqviamosal. 

The frontal has its hinder border sinuously cui-ved, and 
throughout the greater part of its length applied to the parietal. 
Its postero-external is closely applied to, and ultimately fuses 
with, the supero-anterioi- angle of the squamosal and the post- 
orbital process of the alisphenoid. In the supiuorbital region it 
turns downwaids and inwards to form a broad overlapping plate 
articulating with the alisphenoid. Fi'om the mid- orbital region 
onward it becomes band-shaped, ultimately being produced into 
an outwardly directed and blunt angle vmderlying the nasal. 

The squamosal is roughly quadrate in form, and has the superior 
anterior angle pioduced into a lingviiform process which overlaps 
the parietal and alisphenoid. Its antero- ventral angle is produced 
into a small squamosal pi'ominence, the undei- suiface of which 
affords the articular surface foi' the squamosal head of the quadi'ate. 
The postero- ventral angle is obliquely truncated, and foi'ms the 
anterior segment of the rim of the tympanic cavity. The supei'o- 
posteiior angle is produced into a slight point which is wedged 
in between the paiietal and lateral occipital. The hinder boi'der 
of the squamosal serves to cover in the recessus tymjKinicus 
sujjerior, which is foi'med by the absorption of the diploe of the 
lateral occipital. 


Internally the aquiiniosal ;i])peai-s to l)e visi1)le only in Cacaliis 
canoras and (■entro^nis, where it appears as a small triangular 
plate wedged in between the probtic, parietal, and alisphenoid. 

The nasal, judging from the scanty material at my disposal, 
difiei's slightly in form in different geneivi, inasnnich as in Cucalus 
and Centrojyas it is not sufficiently large caudad to cover the 
horizontal plate of the mesethmoid. In Centro])us the hinder 
bordei- is produced into a shai-p angle, and is I'ounded in Cuculus 
and GeoGOCcyx. The nostril is holorhinal. 

The lachrymal offei'S no special featui'es for eounnent. 

The prema.clUa is fully ossified only in the skull of Ceococcyx. 
Its nasal processes are long and slender, and the median suture 
dividing them extends far beyond the level of the anterior border 
of the nasal fossa. The maxillary and palatine processes lie close 
together and parallel with one another. 

The maxilla in Qeococcyx is elongated and triaiigulai- in foi-m, 
and sends backwai'ds from its postero-extei'ual angle a long 
slender bai- to join the jugal and <jua<li'ato-jugal. The Ixxly of 
the maxilla is slightly s[)ongy and has the postero-internal angle 
})roduced backwards and inwards into a maxillo-palatine process. 
The botly of the maxilla lodges a Ijai-ely pei-ceptible anti'um. 

The qaadrato-jugal is long and slender in Qeococcyx ; the re- 
lations between the jugal and maxilla cannot be made out, the 
skull having become disarticulated. 

The vomer is absent. 

The palatine does not differ apprecial)ly from that of the adult. 

The pterygoid is rod-shaped, and prodiiced f oi'wards into a sharp 
triangular spine (PI. XXII. fig. 10), which articulates with a coi-- 
responding facet in the mesial border of the palatine. Later this 
triangular process becomes segmented off fi'om the main shaft 
to form the hemipterygoid, which ultimately fuses with the 
palatine. At the pi'esent stage this segmentation is faintly indi- 
cated by a shallow fuii-ow on the outei- siuface of the shaft. In 
the adult, whei'e the fusion with the hemipterygoid is complete, 
the palato-pterygoid articulation is formed by the approximation 
of glenoid sui-faces sloping obliquely backwai-ds. 

The dentary and splenial do not differ from those of the adult. 

The cm'onoid, angidare, and supra-angidare are all as yet 

iv. The Vertebral Column. 

The vertebral colunni of the Cuculifoi'mes is not marked by the 
same constancy of character which is so conspicuous in the skull. 
In many respects it recalls that of the Coraciiformes, but even 
among the Cuculi i-elatively wide differences obtain. 

All the presynsacral vertebr:>? are heteroccelous, and all the 
thoi'acic vertebi';>? ai-e fi-ee. 

The cervicals of the smaller Cuculi recall those of Leptosoma. 

The odontoid ligament of the atlas is ossified. The axis vei'tebra 
is veiy short antei'o-posteriorly, and has the neui'al ai'ches produced 

Proc. Zool. Soc— 1903, Vol. I. No. XVIII. 18 

272 MR. W. p. PYCRAFT ON THE [Mar. 17, 

outwards and upwards into a pair of pointed hyperapophyses. 
The neural arches of the vertebrae, from the third to the fifth 
inclusive, are very broad, and with or without spines. From the 
third to the sixth or seventh vertebi'pe large metapophyses occui-, 
those of the fourth, fifth, and sixth being especially lai-ge, pro- 
jecting like buttresses from the base of the anterior zygapophyses. 
The metapophyses of the third vertebra send back each a slender 
bar of bone to join the hyperapophysis. A similar bar of bone 
occurs also from the fourth to the seventh vertebrae, but instead 
of joining the hyperapophysis, fuses with the antero-latei-al angle 
of the shield-shaped expansion formed by the neural arch. The 
succeeding cervicals do not afford any particularly well-marked 
characters in so far as this region is concerned. 

The anterior cervicals of Scythrops and Gucuhos (1 to 7) diflfer 
from those of the Cuckoos just described in that they are 
relatively slightly longer antero-posteriorly, that the bony bar 
from the metapophysis to the hyperapophysis occurs only in the 
3rd vertebra, and in that the shield-like expansion of the neural 
arches is very feebly developed. Short neural spines occur in the 
2nd, 3rd, and 4th vertebrae. 

The 2nd, 3rd, and 4th vertebra; bear hypapophyses ; from the 
5th to the 10th these are replaced by catapophyses, which from 
the 11th vertebra backwards to the last cervico-thoracic are 
replaced again by hypapophyses. The cervico-thoracic vertebrfe 
are peculiar in that they bear more or less well-developed catapo- 
physes in addition to the median hypapojDhysis, The catapophyses 
never coalesce to form a carotid canal. 

The cervico-thoi-acic vertebrae may be two oi- three in number. 
The number of the true cervicals varies, being either 11 or 12. 

The thoi-acic vertebra? are free. The 1st or 1st to 3rd, as in 
Eudyncmiys and Coita for example, bear hypapophyses. Pneu- 
matic foramina pierce the centra of the vertebra beneath the 
transverse processes. 

The neui-al spines of these vertebrae gradually increase in height 
from before backwards, and are more or less quadrate in form. 
Coua, however, appears to form an exception to this rule, the spine 
of the 1st thoracic being almost obsolete, and thus agreeing with 
the cervico-thoracic vertebi-ae ; whilst the sjaines of the 2nd, 3rd, 
and 4th vertebrae are, relatively, as high as in other Cuckoos. 

The last two cervico-thoracic (2nd and 3rd) have apparently 
only recently become separated from the thoracic series, inasmuch 
as the vertebral segments of their respective ribs have undei'gone 
no reduction in length. The ribs of the third pair still i-etain 
their uncinate processes. 

Only one thoracic vei'tebra, in the Cuculi, enters into the 

The synsacrum includes from 10 to 13 vertebrae. The smaller 
nuraber appears to have been due to the reduction, possibly by 
excalation, of the lumbar or lumbo-sacral vertebrae, or of caudal 
vertebrae, as in Geococcyx. 


The most complete synsacrum is that of Coua, and is made up 
as follows :— 1 thoracic, 3 lumbar, 3 lumbo-sacral, 2 sacral, and 
4 caudal, making 13 in all. 

Rhoj^odytes seems to possess the most reduced synsacrum : tiie 
number of thoracic, lumbar, and saci-al is the same as in Coua, but 
the lumbo-sacrals are reduced to 1 and the caudal to 2 : making 
therefore a total of 9 vertebra? as against 13 in CoiM. 

Crotophaga and Geococcyx each have 11 vertebije m this region. 
In the former, the reduction is from the lumbo-sacrals, which are 
only represented by a single vertebra; in the latter there are 
2 lumbo-sacrals, but only 3 caudals. 

In other genera, the number of vertebrfe appears constantly to be 
12, composed as follows :—l thoracic, 4 lumbar, 1 lumbo-sacral, 
2 sacral, and 4 caudal. 

In no ca^se can distinct sacral vertebrfe be made out. ilie 
sjns^cvxm-ioiRhmnjyhococcyx is remarka.ble in that the single lumbo- 
sacral vertebra bears a pair of exceptionally strong ventri-latera 
processes. Yestiges of these appear in Scythrojis, Coica, and 


In Centropus the third and fourth pairs of ventri-lateral processes 
are apparently undergoing coalescence. In some skeletons this 
fusion is complete; in others traces, more or less marked, of the 
original buttresses still remain. 

The last two caudal vertebra? combine to form a well-marked 

r)lanum anale. 

There are 6 or 7 post- synsacrals— free caudal vertebrae. Ut these 
the 4th and 6th bear pointed intercentra, which, however, are 
completely fused with their respective centra. _ 

The total number of vertebra? reaches its highest m Coua 

with 37. . . , T ..• • 1 

The vertebral column of the Musophagi is scarcely distmguisli- 

able from that of the Cuculi. 

The cervical vertebra? closely recall those of Cuculus. Ihey niay 
be distinguished therefrom, however, apart from their greater size, 
in that the 3rd and 4th both send backwards a hnv from the meta- 
to the hyperapophysis, and in that the 7th and 8th send back a 
similar bar fiom the same region to the middle of the neural arch. 
There are only two cervico-thoracics. The vertebra coirespondmg 
to the third cervico-thoracic of certain Cuckoos, e. g. Scythrops, 
becomes in the Musophagi thoracic, being joined to the sternum by 
a sternal rib. The thoracic vertebra? differ from those of the 
Cuckoos, in that the transverse processes send l)aclvwards from their 
postero-external angles a long bony spike to overlap the ti-ansverse 
process of the vertebra next behind it. These connecting-rods 
are either wanting or very feebly developed in the Cuckoos, e. g. 

Sc)/throps. . . 

The cervical vertebra? are 12 in number, the cervico-tlioracic Z, 
thoracic 6, the last being fused with the synsacrum. 

The synsacrum contains 13 vertebra?, and is made up as 
follows: thoracic 1, lumbar 3, lumbo-sacral 3, sacral 2, caudal 4. 


274 MR. W. p. PyCRAFT ON THE [Mar. 17, 

The 1st lumbo-sacral shows vestiges of ventri-latei'al processes. 
Only the 2nd sacral bears ribs. 

There are 8 free caudals, but the 7th is nearly fused with the 
pygostyle. The total number of vertebrae is 41. 

I must here point out that the determination of the sacral 
vertebrae in the skeletons herein described is a purely arbitrary one ; 
they cannot possibly be determined with certainty without ex- 
amination of the sacral nerve-plexus, and this I am unable, at the 
present moment, to make. Thvis, it may be that the plexus will show 
that in some cases what is here described as the last lumbo-sacral 
in reality is the first sacral. If this should be the case then, of 
course, the vertebra herein described as the 2nd sacral would be 
the 1st caudal. 

V. The Kibs. 

The cervical ribs, in the Cuculi, form a series of very broad pleur- 
apophyseal lamellae enclosing the usual vertebrarterial canal. These 
plates extend backwards rather be3^ond the middle of the centrum. 
Seen from the ventral surface, the rib is free for about half its 
length. Certain of the vertebrae in the middle of the cervical 
chain have this lamella pierced by a large fenestra. In Guira, 
Coua, Centropi(,s, the 5th, 6th, and 7th are so distinguished ; in 
Crotophaga the 6th, 7th, and 8th ; in Scythrops the 7th and 8th ; 
in Taccocoua, the 4th, 5th, and 6th. fjuculus, Chrysococcyx, and 
probably other smaller forms, differ from the forms just described 
in that the anterior cervical ribs are comparatively long and 
slender, not lamellate ; whilst from the 7th vertebra backwards 
the ribs are reduced to vestiges. 

The cervico-thoracic ribs number two or three pairs. The third 
pair always bear uncinates. 

There are five pairs of thoracic ribs, though not more than four 
are attached to the sternum. The fifth pair are overlapped by 
the preacetabular ilium, and may be reduced to the merest vestiges, 
e. g. Rhamphococcyx and Coua, or they may be complete, as in Scy- 
throps and Cuculus. They vary somewhat in form. For instance, 
in Coua, Centropus, and Guira the proximal end of the rib is 
extremely broad, and the shaft after leaving the vertebra almost 
at right angles, curves abruptly downwards. In other Cuckoos 
the broadening of the shaft is not conspicuous, and the shaft 
slopes gently downwards and backwards, so that the thorax is 
conspicuously broader at the articu.lation with the sternal ribs 
than above. 

The uncinates are broad and strong, and show a tendency to 
develop a sharply defined postero-inferior angle at the junction 
with the shaft of the rib ; this is especially noticeable in Coua. 

The sternal segments of the 5th jDair of thoracic ribs never reach 
the sternum. They may persist as vestiges, even the thoracic 
segment of the rib reaching the verge of disappearance, as in 
Coua and Rhamphococcyx, or they may be of considerable size, 


and extend more than halfway down the sternal segment of the 
4th rib as in So/throps. 

The sternal ribs of Coua reynaucU are i-elatively longer than in 
other Cuckoos, and this fact appeai-s to be correlated with 
degenerate powei-s of flight. As a residt of the lengthened rib- 
segments and a change in the angle formed 1iy the corpus sterni 
and the vei-tebral column, to be discussed presently, the form of 
the thoracic cavity (lifters conspicuously from that of other Cuckoos 
and recalls features characteristic of " Ratitte." These peculiar 
featui-es ai'e wanting in Coiui ccerulea. 

The cervical ribs of the Musophagi closely resemble those of 
the Cuculi, taking the form of broad pleurapophyseixl lamelh^. 
From the 5th to the 8th these lamell* are fenestrated as in the 
Cuckoos, so that the lamella appears to be joined to the centrum 
by a slender bar of bone, the bar extending to the level of the base 
of the postzygapophysis. There are six pairs of thoracic ribs, five 
of which articulate \Wth the sternum. Tlie 6th pair are long, 
but have lost connection with their sternal segments, which 
remain as small spicules anchylosed Avith the sternal segment of 
the 5th pair. In the Cuckoos, it %vill be remembered,^ there 
are only 5 pairs of thoracic ribs, the 5th pair of which, like the 
6th in the Musophagi, fails to reach the sternum. 

vi. The Sterxum and Shoulder-girdle. 

The sternum of the Cuculiformes recalls that of tlie Coraciai 
rather than that of any other group. 

The sternum of the Cuculi may he readily distinguisheil from 
that of the Musophagi in that it is relativel}^ conspicuously shorter, 
being indeed in many cases as broad as long. Moreover it 
appears to be undergoing yet fui'ther reduction. 

The form of the posterior margin of the corpus sterni and the 
relative development of the sjnna externa present considerable 

With regard to the posterior margin of the sternum, this appeai-s 
to be entire in Cacomantis only. Two extremely thin oval areas 
indicate, however, the presence originally of a pair of posterior 
lateral processes, the space between which and the median border 
of the metasternum has been filled up. 

In Scythrops, Cmidus, Coccystes, G^uira, Oentropus, and Croto-^ 
■phaga the posterior border is interrupted by a single pair of 
notches, the outside of the notch being bounded by the posterior 
lateral process. 

All the other genera appear to have a doubly notched sternum, 
but the character of the notches varies consideral^ly. In Playa, 
for example, the posterior lateral process arises directly beliind the 
articulation of the 4th rib, and extends outwards and backwards 
to terminate in a spatulate extremity some distance cephalad of a 
line passing across the free border of the metasternum. Between 
this process and the metasternum is a i\\A\\\^\t processus inter medius. 

276 ME. W. p. PYCRAFT ON THE [Mar, 17, 

which, like the external process, fails to reach the level of the meta- 
sternal border. In Rhinococcyx, Rhamphococcyx, and Taccocoua 
the posterior lateral process arises, as in Piaya, immediately behind 
the articulation for the 4th rib, but it forms a wider angle with 
the long axis of the sternum, and carries with it the intermediate 
process, so that it looks like a branch thereof. In Coita and Geo- 
Goccyx the posterior lateral process leaves the sternal plate further 
back than in the forms just described. In other respects these 
two sterna are quite distinct. 

In Coua ccerulea the posteidor lateral process is veiy broad, and 
extends backwards to the level of the posterior border of the meta- 
sternum, whilst the intermediate process is short and normally placed. 
Coua reynavdi, with an obviously degenerate sternum, differs from 
C. ccEruha in that the posterior lateral process is slender, and the 
intermediate process shows a tendency to fuse with the base of 
the posterior lateral. In Geococcyx the posterior lateral process 
is long and slender, but fails to reach the level of the free border 
of the metasternum ; whilst the intermediate process lies nearer the 
posterior lateral than the metasternum. Further, the sternum is 
peculiar in the great depth of the fissure on either side of the 
metasternum, giving this a long and narrow shape. 

A conspicuous pointed spina externa is found in Gtdra, Citculus, 
Geococcyx, and Scythrops. It is vestigial in Centropiis. A small 
spina interna occurs in Taccocoua. 

In Rhamphococcyx the spina externa and interna are both 
present, and fused to form a moderately large spina communis. In 
Piaya, Rhinococcyx, and Coua there is a vestigial sp)ina communis. 
In Gacomantis the two spines remain distinct, but are extremely 

Both spina externa and interna are wanting in Crotophaga. 

The carina sterni is relatively largest in Cuculus (text-fig. 43, 
p. 277), Cacotnantis, and Scythrops. Its free ventral border is 
strongly arched, and its anterior border is deeply concave. The 
antero-ventral angle of the keel afifords a firm ai^ticulation for the 
furcula. In Coua, Rha^inpliococcyx, Crotopliaga, and Piaya, the 
anterior border of the keel is very deeply emarginate, causing the 
free border of the keel to project forward in the form of a long 
style. With the dorsal surface of the style the clavicle articulates 
by means of a long hypocleideum. Apparently the length of the 
hypocleideum is directly correlated with the i-etreat of the carina 
caudad. In the Galli this is well seen, the climax being attained 
in OpisthoGomus. 

The depth of the carina is extremely i-educed in Coua reynaudi 
(text-fig. 44, p. 277), so much so indeed that the power of flight 
in this species miist be extremely limited. In other Cuckoos the 
depth of the carina is nearly or quite equal to half the breadth of 
the corpus sterni ; in the species in qxtestion the greatest depth 
of the keel is barely one-fourth the breadth of the sternal plate. 
Correlated with the reduction in the keel and the consequent loss 
of flight, is a marked change in the position of the sternum, which 




appeals to liave been tlii'ust forward by a considerable increase in 
the length of the pcistei-ior sternal ribs. As a consequence, the 
shaft of the coracoid nnd the long axis of the sternum form neai-ly 
a, right angle Avith the vertebral column, and the shaft of the 
scapula lies pai-allel with the vertebral axis, instead of forming an 
oT)lique angle therewith. This forward movement of the sternum, 
fui'thermore, has brouglit the aci-ocoracoid upwards to the level 
of the neui-al crests of the vei-tebrte, whilst the distance between 
the acetabulum and the fi'ee edge of the metasternum has lieen 
enormously increased. In fact, the thoi-ax of tins ])ird appi-oaches 
veiy closely, in the relative position of the sternum, that of the 
" RatitJe."" 

Text-fig. 43. 

Text-fig. 44. 

Text-tig. 43.— Left side view of steniiani and shouklcr-givdle of Cucnli'.s c((iioriis, to 
show tile great size of the carina sterni aud the form of the posterior liorder 
of the sternum. 

Text-iig. ii. — Left side view of sternum and shoulder-girdle of Cona rei/naudi, to 
show the degenerate carina sterni, which is actually more reduced tliau the 
figure indicates. Note also the long liypocleidcum and the donhly notched 
corpus sterni. 

Shortening of the sternum is most marked in Piaija. Zaaclo- 
stomios, and Taecocoiia. 

The sternum of the Musophagi is relatively longer and moi-e 
Cora ciif oi-m than that of the Cuculi, but is at the same time thoroughly 
Cuculine in chai'acter. In both Titracus and Schizorhis, the only 
genera represented in the National Collection, the hinder liorder 
of the sternum is doxibly notched, and the notches are of consider- 
able extent, the outei'most being the deepest. The posterior 
lateral process is slender, and directed straight backwards. The 
intermediate pi-ocess lies midway between the posterior lateral 
process and the metastei-nmn. The outermost notch is about 
twice the depth of the inner. 

The co/yios sterni of Turacus is relatively much shorter antero- 

278 MR. W. p. PYCRAFT OX THE [Mar. 17, 

posteriorly than in Schizorhis. The sternal notches of Ttbracus, 
it is to be noted, though preserving the same relative proportions 
between themselves, are yet relatively shallower than in Schizo- 
rhis with regard to their relation to the corpus sterni. 

There is a well-developed spina externa, but no trace of a sjyioia 
interna. In Schizorhis the spina is fiabellate and projects from a 
rounded base. In Turacus it is quadrate in form. In T. huff'oni 
it projects nearly as far forwards as the antero- ventral angle of 
the carina, and is distinguishable from the anterior border of the 
carina only through the medium of a notch. Were this notch 
filled up, tire spina would disappear and the anterior border of 
the keel would present a vertical face pi-ojecting far beyond the 
coracoid grooves, as in certain Ooraciifoi-m birds, e. g. Cyanops. 

Thecoracoid grooves overlap one another. The dorsal lip is 
extensive. A median notch occupies the place of the spina 
interna, and this is bounded on either side by a conspicuous 
oblong glenoid surface, which articulates with a special facet on 
the coracoid. In the Cuckoos the dorsal lip does not overhang 
the ventral, the coracoid grooves do not overlap nor in some 
cases even reach the middle line, and the oblong articular facet 
on the dorsal lip foi- articulation of the coi'acoid is only slightly 
developed. In the Musophagi the dorsal surface of the base of 
the spina extei-na affords ai'ticulation for the right coracoid ; in 
the Cuckoos this is nevei- the case. 

In the Musophagi the ai-ticular surfaces for the ribs ai-e fairly 
widely spaced, less so in Turacus ; in the Cuckoos these surfaces 
are crowded together. 

The coracoid in the Cuculi is relatively long and slendei-, being 
nearly or quite as long as the sterniim. From the ventral aspect, 
one of the most conspicuous features is the large procoracoid. 
This forms a large oblong shelf pi'ojecting inwards and downwards 
from the shaft, at about the level of a line drawn across the shaft 
from behind the glenoid cavity. The acrocoracoid is lai-ge, and 
not twisted on the shaft so as to conceal the foramen interosseum. 
The processus lateralis is well developed and t\\e foramen sup>ra- 
coracoideum is absent. 

The processus lateralis hasalis may be either bi'oad and quad- 
rate, with its antero- external angle produced forwards into a 
spine, as in Cucuhis, Rhopodytes, Eudynaonys, Taccocoua, Rham- 
phococcyx, Scythrojis, and Guira ; or narrow and directed outM'ards 
and backwards, as in Gentropus, Goua, and Bromococcyx. In 
Taccocoua, and to a less marked extent in Gentropus, the impres- 
sion for the sterno-coracoideus muscle, on the dorsal aspect of 
the coracoid, is bounded in fi'ont by a sharp vertically directed 

The scapula is long and narrow, and differs somewhat in shape 
in the different genera ; but the variations are unimportant, and 
not sufficiently^ large to justify desciiption here. The acromion is 
short and stout, save in Goua reynaudi, wherein it is reduced in 
width to form a somewhat cylindrical process. 


The fitrcula lias long, round, slender limbs, gently arched. 
There is a moderately long and styliform hy})ocleideum in Geo- 
coccyx, Cotia, Rhamiihococcyx^ and Playa, for example. In Croto- 
2iha(ja it is spatulate. In Centropus, Cuculus, Eudynamys, and 
Caconiantis it appeal's to ])e of a degenei-ate spatulate type ; 
whilst in Scythrops the h}-pocleideum appears to be wanting. 

The natui'e of the combination of the elements forming the 
inner wall of t\\Q foramen trlosseuvi — the acromion of the scapula, 
the jirocoracoid, and the free end of the clavicle— may prove, 
when exh:iustively worked out, to have some slight value from a 
systematic standpoint. I propose, howevei-, here to offer only a. 
few I'emai'ks, selecting a few genei'a as examples. 

In Cuculus, tScythrops, Co^ia, the whole free end of the furcula 
passes up cephalad of theaci'omion of the scapula and separates the 
latter fi-om the pi-ocoracoid. In Guira. Geococcyx, Rharaphococcyx, 
RJdnococcyx, Rhopodytes, the acromion tuiiis foi'wards so as nesirly 
to join the procoracoid, and forms an opposing surface to the free 
end of the clavicle, which accordingly tui-ns foi-wards to tei-minate 
in a point wedged in between the acromion and pi'ocoi-acoid. Cro- 
tophaga resembles Guira in this i-espect, but the fi-ee end of the 
clavicle is much broader and the articulation with the procoi-acoid 
moi-e developed. In Centropus the aci-omion and acrocoracoid 
meet and embrace, as in a wedge, the styliform free end of the 
slender fui-cula. The pi'ocoi'acoid, which is lai'ge, does not come 
in contact with the scajnila. 

No two of the genei^a,, however, exactly agxee, but the diffei-ences 
between them ai'e too slight to be desciibed in words. 

The coi-acoid of the Musophagi, though closely resembling that 
of the Cuculi, is yet i-eadily distinguishable thei-efrom by the fact 
that the pi-ocoracoid tui-ns downwards and outwai'ds to fuse with 
the acrocoracoid, thus encii-cling the foramen ti-iosseum with a 
continuous bar of bone. The j^i'ocoracoid is large and passes 
insensibly backwards into the coracoid shaft, and thei-e is a 
supracoracoid foi-amen. The p'oces.s«.s lateralis is well developed, 
and directed outwards and ujiwai-ds into a point. In the i-egion 
whei-e the pro- and aci-ocoi'acoid fuse thei-e is an elongated facet 
foi- the ai'ticulation of the fui-cula. The procoi-acoid, however, 
appears to take the gi'eater pai-t of the shai'e in furnishing this 
sui-face. On the doi'sal or intei'nal aspect of the coracoid there is 
developed a special articular surface for the dorsal lip of the 
coracoid gi-oove of the sternum. 

The furcula has relatively shorter, broader, and more laterally 
compi-essed limbs than in the Cuculi. A hypocleideum is wanting. 
The dorsal extremity bears a special projecting facet for articu- 
lation with the procoracoid. 

The scapula I'esembles that of the Cuculi, but is relatively 
slightly broader and has a large acromial process. 

The pro- and acrocoracoid meet and fuse one with the other, thus 
excluding the fuicula from pai-ticipatiou in the formation of the 
foramen triosseum. 



[Mar. 17, 

vii. The Pelvic Girdle. 

The pelvic girdle of the Cnculiformes, though recalling in 
certain f'eatui'es that of the Coraciiformes, is yet quite distinct 
therefrom. Within the grovip it presents a comparatively wide 
range of form. 

Among the Ouculi, the least specialised pelves are those of the 
smaller forms, such as Cacomantis (text-fig. 45) and Chrysococcyx. 

In Cacomantis the preacetabular i-egion of the ilia is sepa- 
i-ated by a low swollen ridge formed by the centra of the lumbar 
vertebrae. The dorsal border is nearly straight; the ventral, 
external, border is deeply emarginate ; the anterior border trun- 
cated and rounded, curving outwards to form a hook -shaped 
antero- ventral angle. The postacetabular region of the ilium has 

CD \ SO 


Dorsal aspect of the pelvis of Cacomantis meruUniis, X 5, sliowing the widely 
separated preacetabular ilia, and the broad dorsal plane of the postacetabiilar 

a broad dorsal plane which abuts against the long and slender 
transverse processes of the vertebrae. These last, by the way, are 
sepai'ated by a row of intertransverse sacral foramina, one on 
either side of the column. The dorsal plane of the postacetabular 
ilium is not produced outwards so as to overhang the ilio- ischiadic 
fissure, and its postero- external border is markedly depressed, 
not forming an upturned crest as in forms to be presently 
desci'ibed. That portion of the ischium which, by its upturned 
growth, converts the obturator fissure into a foramen is very 
narrow ; below and behind the foramen the ischium is produced 


backwards into an obliquely sloping bar. The 2^idn.-i is long, 
slender, and much bowed and closely bound to the posterior half 
of the ventral border of the ischium, lieyond which it extends for 
some considerable distance. The obturator foramen is Ijounded 
posteriorly by a broad bar of bone descending fx'om the ischium 
and fusing with the pubis. The pectineal process is re<luced to a 
vestigial condition. 

In the form of the pelvis, Cuculus, Coccijstes, Scijthrops, differ 
V)ut little, save in size, from Cacomantis merulinus. In Scijthrops, 
however, the obturator foramen is not cut off posteriorly by the 
descent of a bony plate from the ischium. In all save Cacomantis 
and Cuculus the intertransvei'se saci'al foramina are more or less 
completely filled up. 

In Mopodytes and Emlynamys the mesial border of the extreme 
anterior end of the preacetabular ilium rises to the level of the 
neural ridge as in Geococcyx (text-tig. 46, p. 282). In Taccocoua this 
elevated region extends somewhat fui-ther loackwards and forms 
a quadrate plate, the free border of which is lightly applied to the 
nem-al crest. In Crotophaga, Bhamphococcyx, Coua, and Piaya 
actual fusion takes place between this plate and the synsacrum. In 
Centropus and Dromococcyx the relationship between the fused 
region of the ilium and the synsacrum has become still further 
modified, so that the extremity of the ilium has acquired i\ T-shaped 
form, the dorsal limb of the cross-piece bridging a widely open 
canalis ilio-lambalis. A similar canal is of course formed in the 
other cases where the ilia meet the neural ci-est of the synsaci'um, 
but it is inconspicuous. 

Further modifications in the form of the pelvis are the 
enormous lateral expansion of the dorsal plane and the short- 
ening of the pubis. In forms like G^-otophaga, Coua, or Ehain- 
phococcyx, the early stages of the first-mentioned modification 
may be studied. Passmg through forms like Centropus and 
Taccocoua, we reach a climax in Geococcyx (text-fig. 46, p. 282). 
Herein the mdth across the dorsal plane equals the length of the 
synsacrum. This outward extension of the postacetab\ilar ilium 
has resulted in the foi-mation of a huge ledge passing far beyond 
the level of the antitrochanter, and finally, at its free edge, 
turning abruptly downwards and then suddenly upwards, back- 
wards, and inwards, like a pair of wings, so that the surface of the 
dorsal plane acquires a peculiar saddle-shaped appearance. Seen 
from the side (text-fig. 47, p. 282) or from below (text-fig. 48, 
p. 282), this overhanging ledge forms an enormous penthouse 
above and slightly beliind the ilio-ischiadic foramen. A similar 
modification obtains elsewhere only among certain Rails. It 
api^ears to be correlated with a terrestrial mode of life, these forms 
dying but little. 

' The shortening of the pubis is most marked in Geococcyx, 
Piaya, Centropus, and Rhinococcyx, and to a less extent in Coua 
and Crotophaga. 

A large pectineal process occurs in Geococcyx, Coua, Centropus, 



[Mar. 17, 

Rhmnphococcyx, Rhinococcyx, and Piaya ; it is small in Rhopodytes 
and TacGOcoua, and vestigial or wanting in other genera. 

The fovea lumbalis is very small, and the fovea ischiadica and 
jnidendalis are confluent. There is a weil-mai'ked iliac recess. 

Text-fig. 46. 

Text-fig. 48. 

Text-fi2-. 47. 

Text-fig. 46. — Dorsal aspect of the pelvis of Geococcyx mexicanws, showing the 
narrow preacetabular ilia, extremely broad postacetahular dorsal plane, and 
large pectineal process. 

Text-fig. 47. — Side view of same, to show the broad ledge-like plate of the post- 
acetabular ilium, the short pubis, and large pectineal process. 

Text-fig. 48. — Ventral aspect of same, showing the great overhanging ledge of the 
dorsal plane. All figures of natural size. 

The pelvis of the Musophagi resembles that of the more 
specialised Ouculi. Though relatively vi^ide, its breadth is due 
rather to the great length of the synsacral transverse processes 
than to the dorsal plane of the postacetabular ilium. The pre- 
acetabular region of the ilium differs conspicuously from that of 
the Cuculi, inasmuch as in this group the doi'sal aspect is so much 
cut avv^ay as to lie far below the level of the synsaci^al crest, save 
where it rises at the extreme antero-doi'sal ansrle. In the Muso- 


phagi, the preiliuin rises upwards and sweeps iuwai'ds in the foi'in 
of a broad veitical l)lade with a gently lounded dorsal border, 
which ultimately meets and fuses with the synsacral crest, termi- 
nating in a truncated anterior bordei- cephalad of the synsacral 
fusion. The cancdls ilio-lumhalis is wide and spacious. The 
dorsal plane of the postacetabular ilium is moderately wide, and 
tapers posteiioi-ly into a blunt, slightly uptui'ned point. Its free 
outer boi-der is somewhat thickened in Turacas to form a broad 
lip with shai'ply defined ujjper and lower edges ; the infei'ior of 
these edges, immediately behind the ilio-ischiadic foramen, is pro- 
duced into a blunt spine. In Schizorhis the free bordei' of the 
doi'sjil plane is shai'ply defined, so that the blunt point caudad of 
the ilio-ischiadic foramen is apparent on the doi\sal aspect of the 
pelvis, instead of below this level. The ilio-ischiadic foramen is 
relatively much smaller in Taracus than in Schizorhis. The ischium 
expands caudad into an extremely broad plate, especially so in 
Titracus. The fi'ce hinder border of this plate is slightly convex. 
In Turacits the obturator foi'anien is cut off" from the fissure of 
that name by a descending plate of bone from the ischium. This 
is not the case in Schizorhis, the foi'amen and the fissure being 
confluent. The pubis is veiy long, attached foi- some distance 
posteriorly to the venti-al boi-dei- of the ischium. The pectineal 
process is lai'ge in both the genei-a in question. 

The fovea lamhalis is larger than in the Cuculi, and the fovea 
ischiadica is distinct from the fovea piidendalis, the latter being 
cut up into separate compai'tments by the synsaci'al venti-i- lateral 
processes. The iliac i-eceas is faiily spacious. 

viii. The Pectoeal Limb. 

The character of the wing is very uniform throughout the 
group. The humerus only is pneumatic. Most neai-ly resem- 
bling that of the Ti-ogons in its genei-al chai'acter, the limb of 
the Cuculifoiiiies may be distinguished by the greatei- size of the 
pectoral crest of the humei-us, which forms a linguifoi-m or even 
triangular jDlate, and by the pi'ominent coUum trochlece, the 
strongly bowed curve of metacarpal III., an<l the absence of a 
backwardly projecting spur on the uppei- ^ of metacai'pal II. 

In the humerus the coi'aco-humeral gi'oove (sidcns transversum) 
is wanting. The crista superior is gently arched, rising from the 
base of the titbercuhom externus and terminating about the upper 
5 of the shaft. In some cases, e. g. CrotopJutya, the pectoi'al crest 
may be moi-e or less triangular instead of i-ounded. There is a 
very small ectepicond^dar tubercle. The pneumatic foramen is 
small ; the indsura capitis sharply defined. The humerus of the 
Musophagi may be distinguished from that of the Cuculi by the 
fact that in the formei- the pi'oximal Ijorder of the crista superioi- 
is long, low, and coneive, the distal boi-dei- shoit and sti-ongly 

In the Cuculi there is considerable variation in the relative 

284 ME. W, p. PYCRAFT ON THE [Mar. 17, 

lengths of the segments of the wing. Thus, in Scythrops, the 
brachiuni and antebrachium are of equal length, whilst the 
manus is only a very little less in length than either of these 
segments. The brachium is longer than the antebrachium, and 
the latter is in turn longer than the hand in Coua, Rhamjohococcyx, 
RhinocoGoyx, Eiidynamys, Guira, and Groto'phaga. The relative 
disproportion between the length of the brachium and ante- 
bi'achiuni is especially marked in Dromococcyx and Piaya. The 
brachium is longer than the antebrachium, but the latter and the 
manus ai'e equal in length in Centropus and Taccocoiia. In Cuculus 
and Cacomantis the brachium and manus are subequal and 
shorter than the antebi-achium. In Coccystes the antebrachium 
and manus are subequal and longer than the brachium. 

The variation in the length of the wing among the Ouculi 
becomes still more forcibly illustrated when compared with the 
hind limb, the difFei'ence apparently varying with the power of 

In Cuculus, Coccyzus, and Cacomantis, for example, the wing 
is considerably longer than the leg. In Chrysococcyx the two 
limbs are subequal. In Geococcyx, Crotopliaga, and Coua the 
wing is much shorter. Taking the combined length of the femur 
and tibio-tarsus of each species as a standard of measurement, it 
will be found that in Crotophaga the wing exceeds this leng-th 
only by the distance from the middle of Ph. I. D. II. to the tip 
of the wing. In Geococcyx the wing is slightly less than the 
length of the standard. Coua has the shortest wing of all, it 
being about \ shorter than the combined length of femur and 

In the Musophagi the brachium is somewhat longer than the 
antebrachium, but the latter is considerably longer than the 
manus. The ulna, as in the Ouculi, bears prominent bosses for 
the attachment of remiges. 

As in the Ouculi, the collum trochlece is very conspicuous, giving 
the distal end of the humei'us an obliquely truncated form. The 
coraco-humeral groove is wanting. The proximal border of the 
crista superiot- is veiy long and concave, the distal border short 
and convex. The ectepicondylar tubercle is very small. 

The Ph. of D. III. in the Ouculi sends outwards from its post- 
axial border a prominent spur, this is wanting in the Musophagi. 

ix. The Pelvic Limb, 

The pelvic limb in the Ouculiformes has, in common with the 
Psittaci and many Ooraciiformes, a zygodactyle pes. The hypo- 
tarsus is complex, and the tibio-tarsus has an ossified extensor 
bridge. In all save Geococcyx the limb is non-pneumatic, and in 
this genus only the femur is pneumatic. 

The pelvic limb of the Ouculiformes has the 2nd and 3rd 
trochleas nearly equal in length, but the latter is conspicuously 
larger in size. The third trochlea is placed comparatively high 


up the shaft, and has the glenoid articulai' suiface t'onspicuousl>' 
laterally compi'essed and twisted, so as to cross the long axis of 
the shaft transversely. This limb may be readily distinguisheil 
from that of the Psittaci, inasmuch as in the latter the trochle;i 
of D. II. stands out at right angles to the tarso-metatarsal shaft, 
whilst the tiochlea foi- D. IV. has lotated so that its articulai- 
sui-face is tui-ned to look diiecth' back^\"a^•ds. Thus, the outei- 
condyle comes to lie next the shaft, and, furthermore, is pi'oduced 
into a hook-shaped pi'ocess of considerable size. 

Amongst the Coraciiformes, probably the pelvic limb of Leyto- 
soma most nearly I'esembles that of the Cuculifoi-uies, liut the 
conspicuously highei- position on the shaft of the tiochlea of D. IV., 
which obtains in this last gioup, is sufficient to prevent confusion. 

The til)ular ridge is confined to the uppei' end of the shaft, and 
increases in depth from above downwards. 

The length of the fibula ma}' vaiy considerably even in diflerent 
species of the same genus. Usually it is greatly reduced, but in 
Centropus toulox(,, for example, it extends considei'ably be}'ond the 
middle of the tibial shaft, whilst in C. madayascariensis it falls 
considei'ably short of this. 

In Crotophaga, Eudyvamys, and Rhopodytes it barely extends 
beyond the level of the tibial fibular ridge. 

The cnemial crests are, as a I'ule, feebly developed, indeed only in 
GeocoGcyx do they attain to any respectalile size. In this species 
the entocnemial crest is more oi' less quadmte in foi'm ; its outer 
border is convex, its infeiior concave. The ectocnemial ci'est is 
triangular in form, and tei'minates in a small pointed process 
immediately in front of the head of the fibula. 

The shaft of the tibia is long, slender, and cylindiical, and bowed 
slightly forwards. In Geococcyx the posterioi- latei'al bordei-s of 
the intei-nal mesotai-sal condyle ai-e pi-oduced backwai'ds and 
upwards to form a i-athei- prominent spui-. 

The popliteal fossa of the femur is obsolete. 

The pelvic limb of the Musophagi is less specialised in some 
respects than that of the Cuculi. This is especially marked in the 
character of the tarso-metatarsus. As in the Cuckoos, the hypo- 
tarsus is complex, and the tarso-metatarsal shaft grooved in front ; 
but the arrangement of the distal trochlea? is of a less specialised 
character, and this fact is especially mai-ked in SchizorJds, the foot 
of which in this I'espect closely resembles that of Leptosoma. The 
foot of Schizorhis, however, may readil)- l)e distinguished from 
that of Leptosoma inasmuch as in the lattei' the trochlea foi' I). IV. 
bears a very sti'ongly-developed outei- lip, \\hich is pi'oduced 
inwards towards Mc. I. In ,Schizorhis this trochlea is almost 
indistinguishable from that of an ordinary eleutherodactyle foot. 
In Turacus, however, the form of the outer tiochlea nearly 
resembles that of the Cuculi, but is less mai-kedly compiessed, 
and looks backwaids and slightl}- inwards, instead of being twisted 
so as to cross the shaft transvei-sely. Moreovei', it is not raised 
high up on the shaft as in the Cuckoos. 

286 MR. w. p. PYCRAFT ON THE [Mar. 17, 

X. Summary. 

The isolated position of the Cuculiformes among the Coracio- 
morpha? is as evident fi'om a stvidy of the osteology of the group 
as fi-om other points of view. Their nearest allies, judged from 
an osteological standpoint, would appear to be the Coraciidse 
{Coraciince and Leptosomatince) and Bucconidfe on the one hand, 
and — moi-e remotely — the Opisthocomi on the other. Their 
relationship to the Psittaci, which is generally agreed upon, 
would, I think, never be suspected from a compaiison of the 
skeletal fi'amewoi-k. This fact is pi'obably to be explained by the 
great amount of specialisation which the Pari'ots have undergone. 
Concerning the association of the Cuculiformes with the 
Bucconidee, a few words of explanation are necessary. Although 
this connection has several times been made by the older orni- 
thologists, Flii-bringer appears to be the only modern systematist 
who, on anatomical grounds, takes a similar view. Gadow (3) 
regards the Bucconidfe as close allies of the Galbulidte, placing the 
two in the same family — Galbulidee. Beddard, on the other hand, 
widely separates these two forms, and I think rightly so. He 
places the Bucconidfe between the Pici and the Bhamphastidse, 
but remarks that this is a family which " is at present little 
known, and whose affinities are therefore doubtful. It is only 
provisionally that I place them in the present position." 

The claims to relationship, of the group now undei- consideration, 
to the Opisthocomi are, on osteological grounds, not at first sight 
very strong. A careful study, howevei', of the skulls of Cuculi 
and Musophagi, and a comparison with Ojnsthocomus, will show 
points of agreement which suggest affinity i-ather than convei-gence. 
The Musophagi most nearly approach Oinsthocomus. The pelvis 
and sternum of Opisthocomus are Cuculine, 

That the Cuculidfe and Musophagidfe are very near allies there 
can be no doubt. Indeed, the relationship between these two is 
the only relationship about which we can speak with any real 

According to the British Museum Catalogue of Birds, vol. xix., 
the Cuculidse (suboi-der Cuculi) embraces 47 genera and includes 
202 species grouped under 6 subfamilies. This scheme is 
the work of Capt. Shelley. Beddard (1) recognises only 3 sub- 
families, a,nd these, I think, will be found to meet all demands. 
Further, it is quite open to question whether even these should 
not be regarded as groups a, h, c, rather than subfamilies. 

Osteologically, the Cuckoos vary far moi-e widely in respect of 
the pelvis and sternum than in any other character. The great 
featvire in the skixll is the uniform plan of the palate. Trusting 
to this alone, one might divide the Cuckoos into two groups — 
Co'ua in the one, and all the rest of the Cuckoos in the other : 
yet the difference in the two palates is tiivial. The only other 
characters offered the systematist to choose from ai-e the form 
of the lachrymal and temporal fosste and the shape of the beak, 


but the latter at any rate is the direct outcome of adaptation. 
Probably the remarkably specialised condition of the pelvis also 
is adaptive. 

In the accompanying table it will be found th;it I have arranged 
the Cuckoos on a plan formeil of a compromise between that of 
Beddard, Imsed on the pterylography and voice-muscles, and one 
based on the osteolog}' and voice- muscles. The two will be found 
to disao-ree in many points ; and it is on this account that 
I have published the results of my comparison, since it will, I 
hope, stimulate further research by dii'ecting i-enewed attention 
to the anatomy of the discordant forms. 

A. Sternum with sinsflo pair of notches ; pectineal process of pelvis vestigial. 

Mu-icle-foniiuiii AXV-f-, except JEudynamys which has the formula 

a. Syrinx tracheo-bronchial. Coccvstes 

Muscle-formula AXY+, except J Cocr'iam' 
E.d,n,a>,,„s which has ABX\ + . "1 js^J'l^^^; 

01^1 ^^ '^^•l^l- IScyihrops. 

h. Syrinx bronchial. /- rt„„*„„7,^„„ 

Muscle-formula ABXr+ . \ ^^'f"^"' 

New World. ^ ^"'"^- 

B. Sternum with single pair of notches ; pectineal process of pelvis large. 

c. Svrinx bronchial. r n . 
Muscle-formula ABXY+. \ Centropns. 
Old World. iBromococcyx. 

C. Sternum with two pairs of notches ; pectineal process large. 

d. Syrinx bronchial. 
Muscle-formula ABXY-I-. 

New World Geococcyx. 

Old World Coua. 

(^ Rhinococcyx. 
„ . . , , . , I Ehampliococcyx. 

e. Syrinx tvacheo-bronchial. Urococci/v 
Muscle-formula ABXY-H, except j Ejiopodl/te's. 

^, .K'fr^'^'\ ."'^"^" '^'^' "^^^ "*■• Zanclo'stomus. 

Old \\ orld. I Taccocoua. 

{Fiaya. (New World.) 

D. Sternum with two pairs of notches ; pectineal process vestigial. 

f. Svrinx tracheo-bronchial. , ^, 
Muscle-formula AXY-f. \ CJirysococ^yx. 
Old World. ^ Cacomanhs. 

Mr. Beddard has sho^vn, in tabular form, the points wherein 
the skulls of the Cuculi and Musophagi differ one from another. 
These ai'e not many, aiad I cannot but think that they are really 
of slight importance. The most striking are the differences 
in the size of tlie antorbital plate, the position of the maxillary 
end of the quadrato-jugal bar relatively to the tomium, and the 
I'elation of the mesial l)oi'ders of the proximal ends of the palatines. 
The desmognathism of the palate seems to me to be precisely 
similar in both foiins. If the skull of Turacus, for example, be 
compared with that of Coua, it will be found that the chief 
difference lies in the moi'e spongy nasal septum of the latter, 

Proc. Zool. Soc— 1903, Yol. I. No. XIX. 19 

288 . MR. w. p. PYCRAFT ON THE [Mar. 17, 

which descends so as to lie between and force apart the maxillo- 
palatine, causing the palate to be indirectly desmognathous. In 
Turacus^ the nasal septum fuses with the dorsal borders of the 
maxillo-palatine, and does not descend between them, but in this 
case also the palate is indirectly desmognathous. In both Coua 
and Tt(,racus, but for the septum, the palate would be schizo- 
gnathous. But the maxillo-palatine region in ScMzorhis, for 
example, is practically identical with that of normal Cuculi. 
Coua has the least specialised palate among the Cuculi ; Turacus 
among the Musophagi. In the more specialised skull of both 
groups indirect desmognathism appears to have taken place by the 
descent of the ventral border of the septum nasi between the 
maxillo-palatine processes ; resulting in the formation of a flat 
palatal surface, or one with a slight median gi-oove. 

The very striking differences in the form of the pelvic girdle 
stand in strong contrast with the uniformity found in the skull. 
How considerable these differences in the girdle are may be seen 
in the figures on pp. 277, 280, and 282. 

Finally, I may remark that a great deal yet remains to be done 
befoi'e our knowledge of the Osteology of this group is complete. 
Of the nestling skeleton of the Musophagi we know nothing, and 
only a very little concerning the early stages of the Cuculi. 

Only about one-third of the total numbei- of genera of Cuculi 
are represented in the Museum Collection of skeletons. These, 
howevei', fortunately, include all the moi'e important forms. 

xi. Key to the Osteology of the Cuculiformes. 
A. Skull. 

Palate desmognathous, and without conspicuous anterior palatal vacuitj- ; nostrils 
holorhinal, imperforate ; basipterygoid processes wanting ; vomer vestigial or 
wanting ; pterygoid free ; lachrymal free ; antorbital plate large ; the postorbital 
process never extending down to the quadrato-jugal bar. 

A. Mesial border of palatines not meeting in middle line, leaving parasphenoidal 

rostrum exposed ; with a large os uncinatum Musophagi. 

B. Mesial border of palatines meeting in middle line below and concealing para- 

sphenoidal rostrum Cucuxi. 

Key to the Genera of Cuculidfe. 

a. Beak long and pointed. 

a'. Lachrymal with massive descending process ; nostrils linear near the middle 
of the tomium ; temporal fossae confined to lateral aspect of skull. 

Geococcyx . 
b'. Lachrymal very small ; temporal fossae meeting one another to form a broad 

sagittal crest SaurotJiera. 

h. Beak with a high median keel ; lachrymal large Crotophaga. 

c. Beak swollen, with an arched culmen. 

c'. Palatines fused in median line posteriorly ' Scythrops. 

d'. Palatines free posteriorly. 

a". Nostrils linear immediately above tomium ; postorbital process of lachry- 
mal large Rliamplwcoccyx. 

b". Nostrils oval immediately above base of tomium, and bounded above or 
confluent with a conspicuous fenestra ; postorbital process of lachrymal 
small Mhinococcyx. 



d. Beak not exceeding the cvauiuin in length, of medium size, and with a more or 
less strongly arched culinen. 
e'. Descending process of lachrymal vestigial or wanting; temporal tossa 
meeting to form a sagittal crest ; nostrils more or less markedly rem form, 
being formed like those of Ilhinococci/x by the confluence of the narial 
aperture with a superior fenestra. 
c". Descending process of lachrymal in form of delicate incm-ved style. 

a'". Sagittal crest reduced to a mere line Ehopodj/tes. 

b"'. Sagittal crest broad Zanclustomus. 

d". Descending process of lachrymal reduced to a stump. 

c". Descending process of lachrymal wanting; antorbital plate slightly 

inflated Taccocona. 

f. Descending process of lachrymal moderately strong, bent outwards and 
backwards upon the supraorbital ])ortion; temporal fosss meeting to fonn 
a broad sagittal crest. 
f". Horizontal process of lachrymal longer than descendmg; pterygoid with 

d> irsal keel ; interorbital fenestra large Gtiira. 

q" . Horizontal process of lachrymal not exceeding descending process, and 
having free orbital border sharply truncated; interorbital fenestra small, 

g'. Descending process of lachr\nnal slender ; temporal fossa; meeting to form a 

broad sagittal crest : interorliital fenestra very large ... Ce)itropus. 
/i'. TemporarfossiE confined to lateral aspect of skull. 

h". Lachrymal moderately large and not distinctly divided into supraorbital 
and descending processes ;' antorbital plate spongy; interorbital fenestra 
large ; maxillo^palatines not meeting in middle line, but palate edged by 

nasal septum Coua. 

i" . Lachrymal moderately large, with slender descending process; nostrils 

cricular, just above base of tomium Coccystes. 

J". Lachrymal vestigial ; mandible with broad flange at gape. 

c'". Temporal fossai present Cttculus. 

(i'". Temporal fossa; obsolete Cacomantis. 

B. Vertebrae \ 

All the presynsacral vertebra; are free and heterocoelous ; neural arch of atlas 
broad ; hypapophvses blade-shaped, not extending beyond 2nd thoracic vertebra. 
Axis vertebra not conspicuously shoi'tened antero-posteriorly, and not having 
conspicuously upturned hyperapophyses. 
A Lateral borders of thoracic vertebraj terminating in a sharp backwardly directed 

spike MusoPHAGi. 

B. Lateral borders of thoracic vertebrae not produced into spikes. CuccLi. 

C. Pectoral Girdle and Sterxum. 

Coracoid with a well-developed procoracoid; clefts of the posterior notches of 

sternum never extending as far forwards as the level of tlie articulation ot the ribs; 

furcula large. 

A Procoracoid fused with acrocoracoid ; furcula with a well-marked facet for 
articulation with the fused pro- and acrocoracoids ; anterior border ot carina 
sterni continued forwards to level of spina e.xterna M usopuagi. 

B Procoracoid not fused wit)\ acrocoracoid ; furcula without articular facet for 
acrocoracoid ; anterior border of carina sterna not continued forwards as tar as 
free end of spina externa CocuLi. 

1 The presynsacral vertebrx .A' the Cuculiformes are scarcely to be distinguished 
from those of the CoraciituB and Leptosomin<e. 


290 ■ MR. AY, P. PYCRAPT ON THE [Mar. 17^ 

Key to the Genera of Cuculidaj. 

a. Sternal plate with spina communis. 

a'. Posterior border with two pairs of notches. 
a". Hj-pocleideum styliform, free end onlj- overlapped by carina sterni. 

Rlia ni'pliococcyx. 
h". Hypocleideum styliform, overlapped by carina sterni. < ^ tinoeoccyx. 

c". Hypocleidemn oval, overlapped by carina sterni < ■pl°^'^ '•'-' ^*" 

h'. Posterior border with single pair of notches Cocetjstes. 

h. Sternal plate with spina externa only. 
c'. Posterior border of sternal plate with two pairs of notches. 

d". Hypocleideum underlapped b}' carina sterni Zanclostomus. 

e". Hypocleideum with free end onlj' underlapped by carina. Geococcyx. 
d' . Posterior border of sternum with a single pair of notches. 

f". Hypocleideum obsolete ; procoracoid small, affording articulation for 

a'" . Furcula large and stronglj' arched Scythrops. 

V". Furcula slender and slightly arched. 

a*. Posterior lateral process broad Tludynamys. 

b*. Posterior lateral process narrow Ghiira. 

g" . Hypocleideum obsolete ; procoracoid large, not affording articulation for 
scapula Centropus. 

c. Sternal plate with spina interna onlj' Taccocoua. 

d. Sternal plate without spines {nTomoToccyx. 

e. Sternal plate with spina externa and interna distinct. 

h" . Posterior border of sternum with a single pair of notches. 

i". Posterior border of sternum with two pairs of notches . Chrysococcyx. 

D. Pelvic Girdle. 

k. Preacetabular ilium with stronglj^ arched border meeting in the median line abova 
the synsacral spines to form a large canalis ilio-lumhalis . . . Musophagi. 

B. Preacetabular ilia \^'ith dorsal border never meeting in median line above syn- 
sacral spine CtrcuLi. 

Key to the Genera of Ouculidfe. 
a. Presynsacral neural arches broad and without neural spines ; ilia widely 

separated. _ i Cacomantis. 

a. With large intertransverse sacral toramma < n ,„.,i „ 

. No intertransverse sacral loranuna. 

a". Length not exceeding 1 inch Chrysococcyx. 

h". Length not less than 2h inches Scythrops. 

h. Presynsacral neural arches bearing a high median ridge; no intertransverse 
sacral foramina. 
c'. Pectineal process vestigial. 

c". Antero-dorsal angles of pre-ilium not meeting in mid-dorsal line ; pubis 

veiy long JEudynamys. 

d". Antero-dorsal angles of pre-ilium meeting in middle r n^„4.„^i „ 

line; pubis short {Guira! ^''' 

d . Pectineal process large. ^ 

e". Iscliio-pubic fissure slit-shaped and very narrow. 
a'" . Antero-dorsal angles of pre-ilium produced into ^ (Je^^frnvus 

narrow bars meeting in middle line , ^ Dromococ'cyx. 

V". Antero-dorsal angle of pre-ilia produced inwards to form a quadrate 
plate embracing neural spines; postacetabular ilia produced into an 

enormous saddle-shaped plate Geococcyx. 

f". Ischio-pubic fissure extremely reduced. 

c'" . Antero-dorsal angle of pre-ilia produced into a quadrate plate with con- 
cave superior border just reaching neural spine ... Taccocoua. 
d'". Antero-dorsal angle of ilia produced into rounded ^ -oi, ■,,„„„„„,,,, 

angles, not reaching the neural spines | Blwpodytes. 

e'". Ischio-pubic fissure veiy wide, closed posteriorly... Rhampliococcyx. 



(1) Beddard, F. E.— Classification of Birds. 1898. 

(2) FiJRBRiXGER, jVI.— Vei-gleich. Anat. d. Bnistscluilterapparates. 

Theil V. : Vogel. 1902. 

(3) Gadow, H.— Bronn's Thier- Reich. B.l. vi. Vogel. Syst. Tlieil. 


(4) Parker, W. K.— Tmns. Linn. Soc, 2n(l ser. Zool. vol. i., 1879. 


Fi"' 1 Ventral aspect of the stuU of Coua reynaiuli, showing the most primitive 

'" ' fomi ot the indirt'ctly desmoirnathxis skull of the Cuculida'. Herein the 

inuxillo-palatines do not meet in the middle line, but the palate is bridged 

by the swollen nasal septum. Basipterygoid processes are wanting. Ihere 

is no voiuei'. , ^ . , • ^^ • 3- J.^ ^ 

Fi"- " Ventral aspect of the skull of Tiiracus htiffom, showing the indirectly desmo- 

"■ "" >'nathous t vpe of palate of the Musophagidic. The maxiUo-palatines do not 

meet in the middle line, l)ut the palate is bridged b}' the fusion ot the yenti-al 

border of the nasal septum with the dorsal aspect of the maxillo-palatmes. 

The vomer is wantins, but present iu a vestigial condition in Schizo7-his. 

Note the appearance'of the parasphenoid between the palatines. 

Fi"- 3. Dorsal aspect of the skull of Cuua reynaudi, showing the absence ot a nasal 

"' ' hin«-e and the ceiural character of the roof of the skull. 
Fi- 4. Dorsal aspect of "the skull of Turacus leucotis, showing the nasal hinge and 

""' " the o-eneral conformation of the roof of the skull. 
Fig. 0. Dorsal aspect of the skull of SaurotJiera vetula, showing the long upper jaw 

and the large temporal fossae. , ■ ^^ e e >.\ 

Fi" 6 Lateral aspect of the skull of SchisorJiis zonura, showing the form ot the 

°' " narial apertures, the os imcinatnm and the general form of the skull. 
Fi"- 7. Lateral aspect of' the skull of RJiinococcijx calorhi/nchus, to sliow the form 

''' ' of the narial aperture and the ■reneral form of the cranium. 
Fig. 8. Lateral aspect of the skull of a nestling Centropus, showing the arrangement 
of the bones. Outer view. , ,, , • ^, * 

Fi"-. 9. Lateral aspect of the inner view of the same skull, showing the separate 

arrans;ement of the bones. ,. ^ x u ^i, 

Fi"-. 10. The distal end of the pterygoid of the nestlmg Centropus, to show the 

^Explanation of Letters. 

a?<i.=alisphenoid. ».7j.=nasal hinge. 

a.r.. = antorbital plate. qp.=opisthotie. 

apv =anterior palatine vacuity. o.m.=os uncinatum. 

Cc.=basioccii.ital. _^j.=parietal. ;p.a.=palatine. 

c = occipital condyle. ^aj'. = parasphenoidal rostrum. 

ep.o.=epiotic. ^.o.i).=postorbital process. 

e^.o.=exoccipital. ^7-o. = prootic. 

/:/•. = tloccular fossa. p^.=])terygoid. 

fr. = frontal. 'i- = quadrate, 

.■ny =bpminterv!roid. s.e.=sagittal crest. 

A.««.=hemiptervgoid. s.e.=sagittal crest. 

?.=lachrvmal. s.o.=supraoccipital. 

meatus intevnus. s(/.= squamosal, 

mesethmoid. ^/. = temporal fossa. 

««. = nasal. 

292 MR. H. SCHERREN ON [-^W- ^l^ 

April 21, 1903. 

Dr. Henry Woodward, F.R.S., Vice-President. 
in the Chair. 

The Secretary read the following report on the additions made 
to the Society's Menagerie in March 1903 : — 

Tbe registered additions to the Society's Menagerie during the 
month of March were 67 in number. Of these 14 were acquired 
by presentation, 43 were received on deposit, and 10 in exchange. 
The total number of departures during the same period, by death 
and removals, was 89. 

The following papers were read : — 

1. Linnseus and Hunter on Feather-Tracts. 
By Heney Scherren, F.Z.S. 

[Received March 12, 1903.] 

(Text-figure 49.) 

The credit of using the feather-tracts of birds as a means of 
classification belongs undoubtedly to Mtzsch, whose results, edited 
after his death, by Buraieister, were published at Halle in 1840 
under the title ' Pterylographie'. An English edition, tiunslated 
by W. S. Dallas and edited by Dr. Sclater, was brought out by 
the Ray Society in 1867. Pterylosis, or the distribution of these 
feather-tracts, is, according to Prof. Newton, " of prime taxonomic 
importance in Ornithology, though more in the investigation of 
small than of lai'ge groups." This also seems to have been the 
opinion of ISTitzsch himself, who, however, was not aware that 
anything at all had been done even in noting the existence of 
such tracts and of the featherless spaces which he called apteria. 
In his Introduction he says : — 

I maj% therefore, flatter myself with the hope of awakening the interest 
of naturalists by the announcement of my new results, and, bj' the enumera- 
tion and detailed description of the feathered regions of the bodies of birds 
to which I give the name of feather-tracts {joterylce, Federnjlitren), of 
proving that these, new and surprising as they may appear to many on the 
first glance at my figures, really furnish equally significant and important 
characters for the certain and natural discrimination of the families of 

Professor Newton (' Diet. Birds,' Introd. p. 63, note 1) says that 
the only men before Nitzsch's time who seem to have noticed 
feather-tracts wei-e the gieat John Hunter and the accurate 


Macartney. The observations of tlie latter were published in 1819 
(Rees's Cyclopaedia, article Feathers) : — 

Although the common feathers cover the whole body, thej- do not 
gi-ow from every part of the skin ; they ave thickest upon the shoulders and 
loins, along the underpart of the neck and breast, and do not exist upon 
the lateral lines of the neck or breast, or about the un^bilicus. This 
arrangement, and their being directed downwards and backwards, allows 
them to cover the body more neatly, and to remain unruffled during the 
motions of the bird. 

The observations of Hunter appeared first in Owen's ' Catalogue 
of the College of Surgeons ' (vol. iii. pt. ii. p. 311), dated 1836 : — 

Although the feathers of birds appear to be an entire and uniform 
covering, they do not arise equally from every part of the body, but only 
from such parts of the skin as are least liable to be affected by the motion 
of the contiguous parts, such as the motion of the limbs. 

Hunter, however, seems to have done something more, and to 
have discriminated the feather- tracts, for, on the page quoted 
above, we read : — 

To these groups or thickets of feathers I shall give particular names, 
taken from their situation. 

It is, of course, only a coincidence that Hunter used the terms 
" thickets of feathers," and that Nitzsch, in his Essay, ' Pterylo- 
grnphia? Avium, Pars prior,' 1833, chose a very similar name, 
j)tertjlce, which Prof. Newton has translated " feather-forests." It 
may perhaps be of service to recoi-d the fact. 

If the date of Hunter's observations be taken at about 1785, in 
which yeai- he built his Museum, an eai^lier notice of the feather- 
ti-acts exists by at least twenty years. This occurs in the treatise 
By Linnieus, ' Fundamenta Ornithologica,' presented at Upsala by 
A. P. Bf>?ckmann, May 4, 1765 ( Amcenitates Acad. vii.). Premising 
that the feathers ai-e ai'ranged in the form of a quincunx, the 
author pi'oceeds : — 

Nuda vero cutis est (h. e. pennarum quinciince uon perforata ant tecta) 
utrinque ad colli latei-a, a capite interscapulimn versus, et ab axillis per 
latera pectoris ad iuguina usque, atque per femora postica, remotis iutegu- 
mentis, videnda. 

This is as cleai- as language can make it. If, however, any 
doubt could exist, this would be at once removed ])y reference to 
the plate (see text-fig. 49, p. 294), where, in a schematic bird, the 
contour-feathers are ai-ranged in quincvuices, and there the bare 
space running from the neck to the femur is connected with that on 
the opposite "side by the naked patch in the interscapular region. 
The naked spaces, as figured, would, if looked at from above, forn\ 
a quincunx ; and the text i-ecalls a passage in the ' Gai-den of 
Cyrus,' where Browne writes of the quincuncial arrangement in 
the " feathery plantation about liirds." Although the primary 
reference Ijy Browne is to the papillse in the feather- tracts, I suggest 
(and in this I am glad to have the support of Mr, Southwell) 



[Apr. 21, 

that the passage alluded to shows that Sir Thomas Browne was fully 
aware of the distribution of the feathers in well-defined i-egions, 
and that these difi'ered in position and extent in different birds. 
Moreover, I believe the passage in the ' Amcenitates ' shows 
that Linnteus knew of Browne's work. Besides the reference to 
the quincunx already quoted, there is yet another — " Pennfe qufe 

Text-fig. 49. 

Topographical diagram showing feather-tracts and bare spaces in schematic bird. 
(Reduced from ' Amcenitates Academics,' vii. tab. i. fig. A.) 

(preeter alas et caudam) reliquum corpus servant, in quincicncem 
digestsg sunt " ; and his phrase " mirando nunquam satis artificio, 
quod in Colymbi prfesertim corio, alutaiiorum arte prseparato . . . 
est conspicuum," is strangely reminiscent of the note of the famous 
Norfolk natiu'alist : " Elegantly conspicuous on the inside of the 
stripped skins of the dive-fowl." 

by Capt. H. N. Dunn, 
By Oldfield Thomas, 

2. On some Mammals collected 
R.A.M.C., in the Soudan. 

[Received March 11, 1903.] 

The National Museum owes to Capt. H. N. Dunn a collection 
of Mammals, mostly small, made by him in the Egyptian Soudan 
in the course of 1902. Among these no less than five prove to 
need new names, thus showing how much still remains to be done 


in studying the mammals of this interesting region ; and it is 
hoped that other officers Avill follow Capt. Dunn's example. 

The localities at which collections were made are mostly luther 
to the southward of Khartoum, the majority of them being in 
Kordofan, a province hitherto almost entirely inn'epresented in 
our collections. 

Besides the earlier Awitings of Sundevall ^ and Heuglin - on the 
Mannnals of this region, reference may be made to a paper by 
Mr. de Winton on a collection from Shendy \ to the north of 
Khartoum, to the same author's work ' in conjunction with the late 
Dr. J. Anderson on the Mammals of Egypt generally, and to a 
small papei- ' of my own on Mr. HaAvkei-'s collection fiom the 
Fashoda region of the Nile. 


4. c?- Khartoum. 15 August, 1902. 


65. c?. El Obeid, Kordofan. 22 October, 1902. 
ff. caffer had already been recorded from this region by 
Temminck '\ 

3. Megaderma froxs Geofl'. 

9. Wad Medina, Blue Mle. 21 September, 1902. 

4. Taphozous perforatus Geofl". 

1, 2, 3. c? $ $ . Khartoum. 28 June, 1902. 

5. Canis axthus soudaxicus, subsp. n. 

60. $. El Obeid. 2 October, 1902. (B.M. No. 

119. Habessa Wells, W. Kordofan. 7 December, 1902. 

The eastern i-epresentative of the Senegal C. anthus F. Cuv. 
Closely allied to the typical foi-m, but palei- and with mai-kedly 
smaller teeth. 

General characters as in true C. anthus. Colour rather paler, 
a clear sandy buft". Hairs of back and of tail bi'oadly tipped with 
black. Skull, as may be gathered from the measm'ements below, 
smallei' and moi'e delicate. 

Dimensions of the type : — 

Head and body 650 mm. ; tail 230 ; hind foot (s. u.) 137 ; ear 

Skull: basal length 136; zygomatic breadth 78; length of 

1 " Oin Professor J. Hedenborgs insamlingar at' Dasgdjur i NordiJstra Africa," 
K. Vet.-Ak. Hamll. 1&42, p. 189. 
- Keise N.O.-Afrika, ii., 1877. 
3 Nov. Zool. viii. i>. 397 (1901). 
•• Anderson & de Winton, ' Manim. Eijypt,' 1902. 
* Ann. Mag. N. H. (7) viii. p. 273 (1901). 
^ Fide Anderson, ' Mamm. Ecypt,' p. 102. 

296 MR. OLDFIELD THOMAS ON [-^P^^- 21, 

nasals (diagonally) 49 ; intei'oi'bital breadth 26 ; breadth across 
postoi'bital pi'ocesses 37 ; breadth of biain-case 49*5 ; palate, 
length 72*5 ; breadth between outer corners of p/ 44. 

Teeth: length of p.^ 9'6, of -p.^ (on outer edge) 14*2, of m.^ 11, 
of m.^ and m.^ combined 16*8 ; breadth of m.^ 14, of m." 10. 
(Below), length p.3 87, of p., 10, of m.^ 17-3, m., 8-1. 

JIab. and tt/pe as given above. 

This is the Jackal figured by Cretzschmar ^ from Riippell's 
specimen as C. anthus, but is cleai'ly at least subspecifically 
difiei-ent from that animal. Mr. de Winton has recently shown ^ 
that none of the earlier names of Hemprich and Ehrenberg or 
other authoi'S apply to this form, and I therefore venture to bestow 
one on it. 


40. $ . Khartoum. 4 September, 1902. 

7. VuLPES PALLIDA Oretzschm. 

8. c? (young). Wad Medina, Blue Nile. 18 September, 1902. 


118. d" . Gebel Haraza, W. Kordofan. 6 December, 1902. 

The rediscoveiy of this species is of interest, as there has always 
been some doubt whether it was or was not the same as the more 
northern /. lyhica. It proves to be readily distinguishable by its 
smallei' size, by certain differences in its body pattern, and, 
especially, by the absence of the black ends to the caudal hairs. 

9. DiPODiLLus STiGMONTX Heugl. 

5. $. Khartoum. 19 August, 1902. 

A topotype of the species. As ali'eady noted '\ the specimen in 
the Stuttgart Museum marked Meinones stigmonyx does not agree 
with Heuglin's description of this animal, and is more like that of 
his M. dongolanus. 

It is to be observed that the j)resent Gei-bille and the next one 
are so extremely alike, that it is almost impossible to distinguish 
them apart except by an examination of the soles and skulls. 
There is, however, a darker median area (" Scheitel und Biicken- 
mitte sattei- gefarbt ") in the Dipoclillus not present in the 
Gerbillus, and this confiims my previous allocation of the name, 
which in any case having once been made should be adhered to. 

10. Gerbillus agag, sp. n. 

96. d. Agageh Wells, W. Kordofan. 17 November, 1902. 
(B.M. No. Ty2ye. 

A small species of ti-ue hairy-footed Gerbillus, with compara- 
tively shoi't tail. 

1 Atl. Riipp. Reise Mamm. pi. 17 (1826). 

2 Anderson & de Wiuton, Mamm. Egypt, p. 213 (1902). 

3 Ann. Mag. N. H. (7) viii. p. 276 (1901). 


Size small, and feet .short. General colour above soft sandy 
buff, .slightly lined on the back with the dark tips to the hairs, but 
without an}' marked dai'ker dorsal area. Along the l)ack the bases 
of the hairs are plumbeous, but laterally, still within the .sandy area, 
the hairs are broadly ringed with white sul)terniinally, though this 
colour does not show on the sui-faee. Under surface pui-e sharply 
defined white as usual. Cheeks, a prominent patch aliove and 
behind each eye, and another behind the ear white. Whole of 
fore limb white, hind limb with a s;indy line down its outer side, 
the innei- side and whole of feet snowy white ; palms and soles 
thickly hairy. Tail short for this group, pale .sandy above, 
darkening towards the pencilled end ; white below. 

Skull unfortunately broken in the single .specimen. Molars 
markedly smaller and lighter than in the common Soudanese 

Dimensions of the type : — 

Head and body 87 mm. ; tail 100 ; hind foot (s. u.) 24 ; ear 11. 
Length of upper molar series 3"7. 

Hah. and ti/pe as given above. 

This little Gerbille is distinguished from its neighbour 
G.2iygc('ryus by its smaller size and shoi'tei- tail. Its close resem- 
blance to Dipodillus stigmonyx has ali-eady been noted. 

1 1 . Arvicanthis dujtni, sp. n. 

103. S . Kaga Hills, W. Kordofan (about 120 ndles W. of El 
Obeid). 20 November, 1902. 

" Dug out of I'eddish .sandy ciiltivation soil, from among the 
natives' crops of dukhan." — H. X. D. 

A many-.striped species of the A. harharus group; allied to 
A. zehra, but smaller, paler, and with the light and dai'k stripes less 

Size small, the smallest of the group. General pale ground- 
coloui' buf}', the latei'al dai'ker stiipes Ijrown instead of Ijlack. 
Head coarsely gi-izzled bnfiy and brow^n. Centi-al dorsal stiipe 
beginning on the cro\vn, veiy nai-row, blackish, but not so deep a 
black as in ^. zebra; outside this there are on each side five 
unintei'rupted bufiy stiipes, sepai'ated from each othei- ))y broad 
brown bands, each of which is divided down the centi'e into two 
by an intei-rupted band of light, an ai-rangement essentiall}' as in the 
othei' species. The light spaces aie throughout clear buff, and the 
dark lines brown, a clear butf}' line pa.ssing along below the outer- 
most daik line and edging the pure white of the lielly. \\\A. zebra 
the outer lines at least are white, only those near the spine being 
buffy. Eye-ring buffy. Eai's dull ochraceous, without darker 
marking. Arms and legs pale buffy, becoming white on the 
fingers and toes. Tail well-haired ; dull ochraceous aliove, with a 
naii-ow and inconspicuous mesial line of black ; whitish below. 

Skull conspicuously smaller than in A. zebra and the other 
species of the group, with rather largei' l)ulkv ; incisors naii'ower, 
but molars I'ather largei- in propoition. 


Dimensions of the type : — . 

Head and body 90 mm. ; tail 70 ; hind foot (s. u.) 23 ; ear 14. 

Skull — greatest length 28 ; basilar length 22 ; zygomatic breadth 
13; nasals 10x3*4; interorbital breadth 4-4; breadth of brain - 
case 12 ; palate, length 12*2 ; diastema 6"7 ; palatal foramina 5*6 ; 
leng-th of upper molar series 4*9. 

Type. Old male. B.M. ISTo. One specimen only. 

A Fashoda stiiped rat presented by Mr. K.. M. Hawker in 
1901 being identified as lirvicanthis zebra, the present species 
may be readily distinguished from it by its much smaller size 
and more buffy coloration. 

Capt. Dunn tells me that this I'at was very common in the 
cultivated fields of the natives, burrowing in their crops of dukhan. 
It is a vei'y handsome and distinct species, and I have much 
pleasure in connecting his name with it. 

12. AcoMYS WITHERS Yi de Wint. 
73. S . Katul Hills. 30 October. 

13. Leggada tenella, sp. n. 

7. Old 2 . Roseres, Blue Mle. 14 September, 1902. (B. M. 
No. Type. 

" Found in burrows in cornfield ; had 7 young in the womb." — 
H. lY. D. 

A veiy small species of the L. minutoides group. 

General colour of cheeks and sides a clear sandy or ochraceous 
bufi", with a distinct darker median dorsal area, commencing as a 
narrow well-defined line on the nose, broadening on the crown, 
and extending, though less sharply defined, all down the back, 
and dying away on the rump. Under surface pure sharply 
defined white. No lighter raarkings round eyes. Ears small, 
evenly rounded, grey, their edges faintly whiter ; a large and 
prominent white patch behind and below their posterior bases. 
Fore limbs wholly white ; hind limbs with a narrow line of the 
body-colour continued down on the hindei' side to the heel, other- 
wise white. Tail about as long as the body without the head, 
brown above, inconspicuously lighter below. 

Skull smaller and narrower than in the Cape L. iniinutoides, 
with square and well-defined supraorbital edges. Palatal foramina 
ending level with the anterior fourth of m.\ Posterior palate 
elongated, its hinder edge about equidistant from the last molars 
a.nd the fi'ont of the bullse. 

Dimensions of the type : — 

Head and body 50 mm. ; tail 35 ; hind foot 11"5 ; ear 9. 

Skull: greatest length 17' 2; basilar length 14; zygomatic 
breadth 9 ; nasals, length 6"3 ; interorbital breadth 3 ; brain-case, 
breadth 7*5 ; palate, length 9*3 ; palatal foramina 4*0 ; diastema 
5"0 ; length of upper molar series 3*0. 

Hah. and tyj^e as given above. 


This beautiful little species is readily characterised by its strong 
sandy colour, the marked dorsal darkening, and elongated palate. 

14. Jaculus gordoni, sp. n. 

85. $ . Gebel Agageh, \V. Kordofan. 12 November, 1902. 

104. 105. c? 6 ■ 'K.iiga Hills, W. Kordofan. 20 November, 

106. c?- Gebel urn Durragh, W. Kordofan. 25 November, 

Allied to J. jaculus Linn., l)ut larger and difterently coloured, 
and with lougei- eaivs. 

Size rather greater and build stouter than in J. jaculus. 
General colour above, as compared to the yellowish "bufi"" of 
J. jaculus, darker, and nearly approaching to " vinaceous buft" " 
of Ridgway. Laterally, the dar-k colour seems to pass i-ather 
sooner into' the pure white of the under surface. White markuigs 
more extended than in /. jaculus, the cheek, supraorbital, and 
postauricular white patches all large. White hip-stripe large, 
weakened in colour by a faint bufty or brownish sprinkling. Fore 
limbs wholly white. Hinder aspect of thighs like back. Fine hairs 
of feet silvery white, the terminal half of the long digital hairs 
sandy. Tail of tlie usual pattern, its basal portion isabelline bufty 
above ; black subterminal band rather over an inch in length ; 
white tip :|~| in.; longest hairs at end of tail 16-17 mm. in 

Skull shaped quite as in J. jaculus, but larger and heavier 

Dimensions of the type, measured in the flesh : — 

Head and body 120 mm. ; tail 200 ; hind foot(s.u.) 63 ; ear 25. 

Skull : gi'eatest length in middle line 34 ; basilar length 27-5 ; 
zygomatic breadth 24"; tympanic breadth 24-7 ; length of nasals 
oil outer edge 12*5; interorbital breadth 12-8; interparietal 
5-7 X 8-7; palate, length 17-3; palatal foramina 4*7; diastema 
9-6 ; length of upper molar series 5*1. 

Hah. (of type). Kaga Hills, W. Kordofan. Also occurring at 

Ti/2)e. Old male. B. M. No. Original number 104. 

Tliis Jerboa, which I have named in memory of the famous 
General Gordon, difters decidedly from the ordinary Egyptian 
J. jaculus by its larger size and difi'erent colour. The Museum 
had previously received a specimen of it from Omdurman, 
collected by Mr. W. L. S. Loat, but as that was young and without 
skull, it could not be desciibed. 

The only other species which need be referred to is Dipus 
microtis Reichenow ^ from " Samar, in Nord-ostafrica." That was 
founded on a voung specimen, but the description of its teeth 
shows that it was s'utticiently adult not to be the young of the 
present form, than which it is very much smaller ('' Lauflange 

1 Zool. Aui;. X. p. 369 (1887). 


35 mm."), with shorter ears, and with the remarkable character 
(if not clue to accident) of having no white at the end of its tail. 

15. Lepus ^thiopicus H. & E. 

6. c?. Shendy. 1 September, 1902. d 6 2 2 ■ Agageh, Kordofan. November 1902. 

39. c? . Wad Medina, Bhie^Nile. 

16. Procavia ruficeps H. & E, 

77. c? • Kaga Hills, Kordofan. 2 November, 1902. Agageh Hills. November. 

17. Oryx algazel dammah Cretzschm. 

(5 . Kordofan, 

Practically a topotype of the subspecies. 

When the ' Book of Antelopes ' was published, owing to the 
predilection of the senior author for ''well-established" names, 
the term leucoryx was used for the species to which Lichtenstein 
erroneously transfei'red that name, while the ti'ue leucoryx of the 
Persian Gulf was called by Gray's name heatrix. To put matters 
more in accordance with modern ideas on nomenclature, the 
Scimitar Oryx should bear the name algazel Oken ; but as that 
name was founded on Cuvier's figure of a Senegal specimen, and 
it is practically certain that the Eastern Soudanese foi'm is at 
least subspecifically distinct from the Western, a second name is 
required, and this we find in the " Antilope dammah " of Riippell 
and Cretzschmar. A. dammah was erroneovisly identified by 
Riippell, and following him by Sclater and myself, with the Beisa, 
but by its locality (" die gi-ossen Steppen von Haraza ") is clearly 
shown to be the Scimitar Oiyx. This is fortunate, as the name 
dammah, being earlier than heisa, would have had to be used in 
the latter's place had Riippell's identification been correct, but 
now the Beisa is left with its familiar name unaltei-ed. 

Briefly put, the nomenclature is as follows : — 

Scimitar Oryx. 

Oryx algazel Oken. 

0. leucoryx of authors generally, not of Pallas. 

Western form : — 

Cemas algazel Oken, Lehrb. Nat. iii. pt. ii. p. 741 (1816) ; 
ex L'Algazelle, F. Guv. H. N. Mamm. i. pi. 376 (1819). 

Eastern form : — 

Antilope dammah Cretzschm. Atl. Riipp. Reise, Mamm. 
p. 22 (footnote), 1826. (Haraza, Kordofan.) 

White Oryx. 
Oryx leucoryx Pall. 

Antilope leucoryx Pall. Spic. Zool. xii. p. 17 (1777). 
Oryx heatrix Gray, P. Z. S. 1857, p. 157, and of authors 
generally.^ 2/ 


3 4- 

, y 

\ I 

', Nfc^t 

^'^ P '" a.TTi'.l.^^ 


i .'^ 

'. , /. ;^ 


*^ cc.w. 




4- • 

, /^-?"0 • 

.' •'. •;>• ' •. • < . . 



Bale &, Danielsson L**^ lith. . 



18. Gazella ruficollis H. it; E, 

61. (S ■ Gebel Tneis, 90 miles E. of Omrlin-mrtn. 9 October, 

This fine Gazelle is a most acceptable acVlition to tlie National 
Collection, in which the species is still badly i-epresented. 

19. Gazella rufifrons Gray. 

d' 2. Agageh Wells, Kordofan, November 1902. 
Capt. Dunn tells me that this Gazelle does not range beyond 
about 50 miles to the northward of El Obeid. 

3. On a Collection of Tnrbellaria Polycladida from the Straits 
of Malacca. (Skeat Expedition," lb99-1900.) By F. F. 
Laidlaw, B.A. Cantab., Assistant Lecturer and Demon- 
strator in tlie Owens College.^ 

[Received March 19, 1903.] 

- (Plate XXIII. & Text-figures 50-56.) 

The collection described below was made entirely by Mr. Evans 
on the shores of a small islet called Pulau Bidan, a few miles noith 
of Penang, It proves a most intei'esting one, and includes, so far 
as I can discover, only one pi'eviously known species, Thysanozoon 
auropunctatuvi Coll. 

As in describing Mr. Gardiner's collection from the Maldives 
[4], I have to note the scarcity or absence of the Eurylejotidae, 
i-epiesented doubtfully only by a much damaged fi-agment in 
Mr. Evans's seiies. 

The only previous record of species from the shores of the 
Malay Peninsula that I have been able to discover was made by 
Collingwood, who desci'ibed and figured the following species from 
Singapoie : — 

Thysanozoon allmani. 
Proceros hancockianus. 

„ huskii. 
Sphyngiceps lacteus. 

Eurylepta kelaartU. 
Elas'inodes ohtusam. 
Leptoplana aurantiaca. 

Grouping these specimens with those of the Skeat Collection and 
adding Pseudoceros hedfordii fi'om Singapoi'e, kindly given me by 
Mr. Lanchester, we get the following list classified according to 
Lang's system [6] : — 



* Planocera sp. 
1.* Notoplana evansii, sp. n. 

1 Communicated by Dr. S. F. Haemek, F.Z.S. 

2 For explanation of the Plate, see p. 318. 

302 MR. F. F. LAIDLAW O^S THE [-^W- 21, 

Acotylea (con.). 


2.* Semonia penangensis, sp. n. 

3.* Leptoplana malayana, sp. n. 

4. „ ohtusitm (Coll.). 


5.* Bergendalia cmomala. 

6.* Latocestus argus. 




oon alhnani. 






'OS hancockianus (Coll.). 



huskii (CoU.). 



kelaartii (Coll.). 



hedfordii, sp. n. 



collingwoodii, sp. n. 



'OS ? ruhellus, sp. n. 



Asthenoceros woodivorfhi, sp. n. 

Prosthiostomid^ . 


Prosthiostomum 2}cdlidu7n, sp. n. 



aurantiacum (Coll.). 

Species marked with an asterisk were collected by Mr. Evans. 

In this eomniunication I have not ventured to give an account 
of the anatomy of any of the species, contained in the collection, 
which can be regarded as being at all complete. Siich an account 
would have expanded the paper beyond limits reasonable in a 
systematic description. I have attempted only to give such a 
diagnosis of each species as shall render its future identification a 
matter of tolerable certainty, and to call attention to any of the 
more striking features which presented themselves. 


Planocera sp. (Plate XXIII. fig. 1.) 

A fragment of tissue of which I made microscopic sections 
proves to belong to a species of this genus, but being only a 
fragment I will not attempt to describe or name it, though it is 
obviously new. The section passes through the penis and prostate, 
and the spines lining the Ivimen of the f ormei- are of such a remark- 
able character that I venture to figure one of them, in the hope 
that a perfect example may be obtained ere long. The length of 
each spine is roughly "08 mm., its breadth at the base "02 mm. 

ISToTOPLANA, gen. nov. 
NoTOPLANA EVANSii, sp. n. (Plate XXIII. fig. 2.) 
A number of specimens belonging to this species were taken by 


Mr. Evans on the shoi'es of the island of Pulau Bidiin, whei'e it is 
evidently one of the commoner species. 

The body of this worm is rather elongated, rounded in front 
and with a pointed hindei- extremity. The arrangement of its 
eye-spots is shown in the accompanying figure (text-fig. 50). 

Text-fig. 50. 


• • 

Eye-spots of Notoplana evansii. 

Unfoi'tiinately I have no notes as to coloui-, but, judging from the 
spirit-specimens, the ci-eature is probably of a yellowish-grey 
coloui-. with a small niunber of ii-i'egularly scattered black spots. 
The gut-bi'anches aie numei'ous, and thei'e is no anastomosis. 
The dimensions of an avei'age individual ai'e as follows : — 

Length 25 mm. 

Breadth 15 „ 

Tentacles f i-om ant. naargin 5 „ 

^louth-opening ,, 12 ,, 

(5 . Aperture behind mouth 3*5 ,, 

9. ,, behind male 1"5 ,, 

Genital Apparatus (see text-fig. 51, 304). 

The most stiiking featui'es of the male copulatoi-y appai-atus 
ai'e the great length of the antral chambei- and the chambered 
pi'ostate organ, the latter recalling the jii'ostate of certain 
Leptoplanas, e. g. L. alcinoi. From the male aperture the antral 
cavity (text-fig. 51, a.m.) extends forwards as far as the level of the 
hinder end of the pharynx. It has very muscular walls and runs 
sloping in an upward dii-ection. Lang, in his diagram of the 
genital appaiatus of Hoploplana {Planocera insignis gi-oup [6]), 
calls this chamber the penis-sheath (" Penisscheide), but this term 
should, I think, be reserved for the sheath-like folds of the walls 
of the anti-al chamber occurring in Cestoplaaa and in certain 
Cotylean genei-a. 

The lining of the walls of the antiul chamber in the pi'esent 
species consists of a flattened ciliated (?) epithelium ; the muscular 
wall consists of a thick layer of inteilacing circular fibres. 

Proc. Zool. See— 1893, Vol. I. No. XX. 20 



[Apr. 21, 

The penis (jo.) itself is small and projects into the antral chamber 
at the upper anteiioi- end of the latter. 

The penis is a fleshy organ, composed of longitudinal and 
circular fibres which exhibit numerous nuclei (see Plate XXIII. 
fig. 2). It is armed with a chitinous stylet. 

The base of the penis is entei-ed by the short ductus ejaculatorius 
{d.e.) which leads to the prostate region. It turns shortly after 
leaving the penis and runs in a backward direction, so that a 

Text-fig. 51. 


Male organs of Wotoplana evansii. 
Tor explanation of lettering, see p. 318. 

transverse section passing through the penis passed also through 
the prostate (see PI. XXIII. fig. 2 and also text-fig. 51). 

The prostate consists of a number, some nine in aU, of small 
chambers {pr.c.) which lie about the ductus ejaculatorius and 
open into it at their distal ends. The ductus, elsewhere with 
muscular walls, here is lined only with a very flattened epithelium,, 
outside which lies a second layer of similar cells belonging to the 


inner walls of the pi-ostate chambers. Between each of these 
chambers a fine double septum runs out from this inner wall (Plate 
XXIII. fig. 2, s.). The prostate cells stain very feebly, and the 
lumen of each chamber is well defined, and contains in most cases 
a cei'tain amount of secreted mattei-. Outside tlie chambers is a 
fairly thick layei' of circular muscle-fibres, and, l>e}-ond these, an 
ill-defined layer of cells which appear to be glandular. Here 
and thei-e are faint indications of pi-ocesses from these outer 
glandulai- cells, piercing the muscle-layer of the prostate organ. 
This muscle-layer is also ti-aversed by processes from the septa of the 
chambei-s. Thei-e is no longitudinal muscle-coat to the pi-ostate. 

Towards the level whei-e the prostate-chambers open into the 
ductus ejaculatorius, some of them open into each other, so that 
the number of chambers seen in cross-section is reduced to four 
or five. 

On the proximal, hinder, side of the prostate, the ductus is 
continued back into the vesicula seminalis {v.s.), which is long and 
contorted. It is lined with a ciliated epithelium, and its walls are 
composed of i-egulai-ly airanged cii-culai- muscle-fibres. At its 
exti-eme hindei- end it turns shai'ply forward and ends bUndly. 
Just before it takes this turn forward, the two vasa deferentia 
open into it, one on either side, aftei- piercing thi-ough its 
muscular wall. 

Thei-e is a spacious anti-um femininum suiTOiuided by the large 
shell-glands, the secretions of which it receives. The vagina leaves 
the anti'um dorsally, and i-uns forward for the first part of its 
course thi'ough the shell-glands, the secretion of which it also 
I'eceives. It then turns first doi'salwards and then backwards, 
acquiring in its course a fine wail of circular muscle-fibi'es. After 
I'unning back for a short distance, it receives the shoit common 
duct from the two uteri. It then continues to I'un back as fai' as 
the level of the antrum femininum, where it ends blindl}-. 

It is evident that this species is closely allied to von Plehn's 
Playiotata pi-omiscaa [7] ; in fact, the terminal parts of the male 
apparatus in the two species have almost identically the same 

It difiers from von Plehn's species in having the phar3'nx of the 
noi-mal t}^e, not elongated transversely. This character is of 
sufficient importance to justify the placing of these species in two 
sepai'ate genera. In addition Notojjlana is characterised by 
differences in the female apparatus, and by the gi-eat length of 
the antral chamber of the male oi'gan. 

The genus yotoplana may be defined, then, as follows : — 

A Planoceroid genus ivlth styliform penis and unthout a bursa 
copulatrix. The male antral chamber is very long ; there is a 
complicated prostatic organ consisting of several chambers lying 
aroiond the ductus ejaculatorius in front of the vesicula seminalis. 
The latter is long and twisted. Body rather elongate, unthout 
marginal eyes ; mouth- ojjening rather behind the middle ; pharynx 
normal ; gut-branches mtmerous, ivithout anastomosis. 




[Apr. 21, 

fi'OBi the anterior end. 

Family LeptoplaniDvE. 
Leptoplana malayana, sp. n. (Plate XXIII. fig. 3.) 

A number of specimens ; some, preserved in formol, are of a 
uniform greyish-brown. 

Length, about 35 mm. 

Eyes, about 8 ,, 

Mouth-opening, abou,t 15 ,, ,, ,, 

c? aperture, about ... 10 ,, from the hinder end. 

$ ,, ,, ... 1 ,, from male. 

The arrangement of the eye-spots is that usual in the genus : 
there are two small clusters of " tentacle- eyes " about 1*5 mm. 
apart, and in front of these are a few scattei'ed spots on either side 
of the middle line. 

This species is closely allied to L. paciUcola von Plehn [7]. 

Genital Apparatus (see text-fig. 52). 

The vasa deferentia (v.cZ.) open into the proximal (anterior) end 
of the vesicula seminalis (v.s.) ; the latter, of considerable length 
and much twisted, is lined with ciliated epithelium, and has thick 

Text-fis. 52. 



ace. ves. 

Genital apparatus of Leptoplana malayana. 

a.f., antrum femininum ; acc.ves., accessory vesicle ;, sliell-glands ; 

u.t., uterine duct ; v.d., vasa deferentia. For other letters see p. 318. 

muscular walls made up of circular fibres. At its distal end the 
vesicula narrows into the ductus ejaculatorius (d.e.), which is 
shorter, but, like the vesicula, much twisted. Its walls are like 
those of the vesicula, but it is much narrower and has a less well- 
developed muscle-layer. The penis is small, fleshy, and unarmed, 
and projects into the large antral chamber, which has thick mus- 
cular walls and extends vertically upwards almost to the level of 
the dorsal body-wall muscles. (See Plate XXIII. fig. 3.) 

The female opening lies close behind the male, and leads into a 
moderately large antral chamber {a.f.) with musculai- walls. Into 
this the vagina opens from above. This latter organ is a long and 
tortuous duct of varying diameter ; the shell-glands ( open 
into it through nearly its whole length. At its hinder end, near 


the point wJiere tlie uteri open into it, the muscuhir walls, which 
are elsewhere feebly developed along its course, become thicker, 
but not very mnrkedly so (cf. L. caUfoniica von Plehn). The 
two uteri i^at.) unite as they enter it from below, and behind them 
there is a veiy small accessoiy vesicle (acc.ves.) with muscular 

The genus Leptoplana, as at present constituted, contains a 
consideiable number of species, many of them only i-eferred by 
Lang with doubt to the genus. In the majority of the species, 
the anatomy of which has been investigated by means of sei'ial 
sections, the genital ajipai-atus pi-esents the following chai-acters : — 

The penis is directed backwai-ds. The vasa deferentia open 
into a muscular vesicula seminalis ; from this the ductus ejacula- 
torius I'uns backwai-ds to open into a prostate organ, -which maybe 
chambered ; le;xving this, the duct runs back into the penis, 
which may be armed with a chitinous stylet. 

The vagina runs back from the antrum femininum, receives the 
secretion of the shell-glands, and further l)ack the two utei-i open 
into it ventrally, usually by a connnon duct. An accessory vesicle 
is generally present. The most familiar species of the genus, L. 
tremellaris, differs from the majoiity of species in the structure of 
its male apparatus sufficiently to permit us to put it on one side 
to foi-m of itself a section of the genus fui-thei- characterised hy 
the possession of a ventral sucker between the genital openings. 
Anothei' species, L. suhviridis von Plehn [8] ( = X. 2^<^''''^dalis mihi 
[4]), approaches Biscocelis tigrina in the structure of its female 
oi-gans, and accoixlingly we may put this species also in a section 

The remauiing species I have attempted to classify beloA\-, so f;ir 
as is at present possilale, according to the sti-ucture of the penis 
and prostate. I trust that I will not )>e thought to have laid too 
much stress on the structui'e of these organs in dealing with this or 
other genera. It seems to me that, of all the characters that 
present themselves for classifying this order of Turbellaria, these 
ai-e the most useful, and that they are as reliable as any of the 
other chai'actei-s employed foi' this purpose, such :is structui-e of 
the pharynx, number and arrangement of eye-spots, presence or 
absence of tentacles, &c. &c. 

I propose, then, to group the species in two sections. A, and B, 
so fai- as oui- knowledge of theii- anatomy pei-mits : — 

A. Penis provided with a. stylet. 

(a) Prostate complicated by i-adially arrangetl "Drusen- 

1. L. kukenthali v. Plehn. Spitzbergen [7]. 

(6) Prostate dividerl into chambers lying parallel to the 
ductus ejaculatorius. 

2. L. vitrea L:nig. Mediterranean [61. 

3. L. alcinoi Schmidt. „ [61. 

308 MR. F. F. LAIDLAW ON THE [Api'. 21, 

(c) Prostate not chambered. 

4. L. pcmamensis v. Plehn. Gulf of Panama [7]. 

5. L. californica v. Plehn. California, Chatham Is. [11] 

6. L. nationalis v. Plehn. Ascension Is. [8]. 

The following also belong to section A, but their position in 
the section cannot be determined without information as to the 
prostate characters. 

L. variabilis Giard. 1 

L. ellipsoicles Yerrill. I New England [13]. 

L. virilis Yerrill. J 

L. drmhachensis Oersted. Norway [6]. 

L.fallax (de Quatrefages) [6] should perhaps form a separate 
section of the genus on account of the great length of its stylet. 
It occurs on the south side of the English Channel. 

B. Penis unarmed. 

a. Prostate chambered. 

7. L. chierchcB von Plehn. Callao [7]. 

/3. Prostate not chambered. 

8. L. pallida (de Quatrefages). Mediterranean [6]. 

9. sp. (unnamed). Maldives [4]. 
y. No definite prostate. 

10. L. jmcijicola von Plehn. Valparaiso [7]. 

11. Z. malayana, sp. n. Straits of Malacca. 
Belonging to this section but of uncertain position in it is 

L. ancjtbsta Verrill [13] from New England. L. lacteoalba Yerrill 
[14] is insufficiently characterised, but said to be like L. pallida. 

Semonia penangensis, sp. n. 

Two specimens from Pulau Bidan. No note available as to 
colour. This interesting species agrees fairly closely with the 
only other species of the genus at present known, viz., S. inacu- 
lata von Plehn, from Java [7]. It is, however, readily distinguished 
from that species by its external characters, the arrangement of 
the eye-spots being obviously very diiferent and showing some 
approximation to the condition found in Discocelis tigrina (see 
text-fig. 53, p. 309). 

The dimensions of the larger specimen are as follows : — 

Total length, about 22 mm. 

Breadth, about 11 „ 

Mouth-opening, about ... 9 „ from the hind end. 

Genital aperture 1 „ behind mouth-opening. 

Brain, about 4 „ from the anterior margin. 

The marginal eyes extend back for abou:t 8 mm. on either side. 
The arrangement of the brain and tentacle-eyes is shown in 
text-fig. 53, p. 309. 

An important distinction between this species and Semonia 
macidata is afibrded by the fact that in the latter the testes and 


ovaries lie on the dorsal side of the Ijody (see von Plehn [7]), 
whilst in the present species the position of the testes is normal. 
i. e. on the ventral side. 

In one or two respects this species approaches the closely allied 
Discocelis tiyrina, and is pei'haps intermediate between that species 
and *S'. maculaia. But the latter s})ecies is also certainly closely 
jillied to Discocelis, and I am rather surprised to find that von 
Plehn has not instituted any compaiison between the two genera. 
/S. penanyensis difiers fi'om S. maculaia, and approaches Discocelis 
tigrina, not only in the features already mentioned, viz., the 
arrangement of the eye-spots and the position of the testes, bnt 
also in that the utei'i unite to enter the vagina by a shoi't common 
duct, and in liaving a veiy V)hnit, almost square penis. 

Text-fie-. 53. 


" Braiii-eyes " of Semonia penangensis. 

It difiers from Discocelis, and resembles S. maculaia, in being 
without the characteristic large prostatic cells which occur in the 
epithelium of the penis and of the antrum, and without the 
remarkable paired structiu'es which run foi'wai-d fi'om the accessory 
vesicle oi Discocelis tigrina. Further, like S. maculata, it possesses 
a definite vesicula seminalis, which appears to be absent in 

Lastly, *S'. penangensis is without the cuiious glandular vesicles 
found along the anterioi- ends of the vasa deferentia in S. maculaia ; 
and the vagina is prolonged, beyond the point whei'e it I'eceives 
the openings of the \iteri, into a veiy small accessory vesicle — so 
small, in fact, that it almost escaped observation. This may be 
due to the fact that the only specimen available for section-cutting 
was not quite mature; oi- moi-e probably, since the uteri were full 
of apparently ripe eggs, that tliis organ is undei'going degeneration. 

It is obvious that the distinction between Discocelis and Semonia 
is a very slender one, but still, I think, sufiicient to wari-ant the 
retention of the latter as a valid genus. The most important 
chai-acters separating the two genera are the absence of a vesicida 
seminalis in Discocelis, and the curious hoi-seshoe shape of the 
accessory vesicle in that genus. 

310 MR. F, F. LAIDLAW ON THE [Apr. 21, 

Family Oryptocelidid^. 

Bergendalia anomala, gen. et sp. nov. (Plate XXIII. figs. 4, 
5, 6, & 8.) 

A most remarkable and interesting form, probably allied to 
the anomalous genera Gryptocelides and Polypostia, desci"ibed by 
Bergendal [1], and provisionally referred to the same family with 

The structvire of the female tei-minal ducts is, so far as I know, 
unique, and approached only b}-- the sj)ecies referred to Trigono- 
poms and Polyporus (see von Plehn [11]). 

Only a single specimen was obtained. It is rather a large 
form, with a total length of about 60 mm. and breadth about 
27 mm. The mouth- oj)ening is some 15 mm. from the hind end, 
and the genital pores lie halfway between these two points. 

The margin of the body is complete, surrouiaded by a continuous 
rather dense row of eye-spots. Thei"e are none of these, apparently, 
over the brain. 

Ooloui- in the spirit-specimen uniformly greyish white (see 
PI. XXIII. fig. 8). The specimen appears to be in an early 
stage of sexual maturity, since no trace of ovaries or testes can be 

The pharynx is large and much folded, the gut-branches are 
numerous and anastomose freely. 

The cells of the epidei-mis are elongated, especially on the dorsal 
surface. True rhabdites are absent, but in place of them the 
epidermal cells are crowded with pseudorhabdites which are of a 
coarsely granular texture, faintly stained and columnar in shape. 
In my sections (stained with Grenadier's htematoxylin) certain 
gland-cells lying within the muscle-layers of the body-wall, 
especially on the ventral side, are deeply stained ; from these cells 
I'un processes Avhich piei'ce the muscle-layers and basement- 
mLembrane, and make their way to the surface through the 
epidei-mal cells. These deeply-lying gland-cells and their processes 
have rather a spongy appearance, due to their preserA'ation not 
being quite perfect. 

Genital Organs (text-fig. 54, p. 311). 

The penis (p.) is a small fleshy organ composed of nucleated 
longitudinal fibres (PL XXIII. fig. 4). Its outer side is lined with 
cells continuous with those lining the antrum masculinum, but 
whereas the latter are ciliated, those coveiing the penis are non- 
ciliated. The base of the penis is pierced by a duct rvmning 
nearly vertically upwai-ds. Immediately after leaving the penis 
this is joined by two small ducts — the vasa deferentia {y.d^ — 
which run forward on either side of the middle line. In one of 
them, at the level of the hinder end of the pharynx, there is a 
slight dilatation containing spei-matozoa. 

The vesicula (j'Jr.) lying above the penis is small. Its wall 


consists of a very tluii l;iyer of circular nmscle-fibres. lineil with 
a cubical epithelium, the nuclei of the cells of which lie close to 
the lumen, and which has rather the appearance of an exhausted 
secretory tissue. This oi'gan I call the prostate (PI. XXIII. fig. 4, 
pro.). About halfway between the penis and fenxale aperture lies 
an oi-gan which beai-s some i-esemblance to the penis already 
described. This organ constitutes one of the most interesting 
features of this anomalous species. It consists of a small pro- 
trusible penis-like organ (7;.?), about one-third the size of that 
first described, lying in a small antrum. It is pierced by a short 
duct, which luns into it from a small non-muscular vesicle which 
lies immediately above it (jjr.f). This small vesicle is of about 
the same size as the prostate. Thei'e is no communication, so far 
as I can discover, between the duct connecting these structures 
and the deferentia (PI. XXIII. fig. 5). 


Geiiitnl apparatus of Bergendalia anomala. 

ace, accessor}' part of vagina ; p. ?. penis-like organ ; pr. ?, prostate-like organ ; 
s., spiral coils of vagina. For other letters, see p. 318. 

I can suggest only two explanations of the presence of this 
second penis-like organ and its accessories. 

Fii-stly, that it may be regarded as a pi-ostatic sti'uctiu'e ^^"hich 
has lost its connection with the penis and developed an intro- 
mittent terminal part of its o^\•n. Such a state of afiairs is found 
in the Ootylean family Diposthiidte ; but in that case both penis 
and pi-ostate open into a common antrum, and there is what 
appears to be a pi'ostate gland in connection with the penis in 
Bergendalia, a fact which makes sti-ongly against this view. 

The second possible explanation is that the oi-gans under dis- 
cussion ai'e the vestiges of a second penis. The fact that they l^ear 
a close resemblance to the functional penis tends to support this 
view, as does also, I think, a comparison with Cryjitocelides of 
Bergendal [1]. 

This is the only other Polyclad described, .so far a.s I know, in 
which two penial organs lie behind each other on the middle line; 
but, according to Bergendal, they lie behind the female aperture, 
and, further, lioth open into a common antrum, whilst in some cases 

312 Ma. F. F. LAIDLAW ON THE [Apr. 21, 

thei'e may be four or even six penes present. In another genus 
presumably allied to this, viz. Polypostia Bei-gendal [1], the penes 
lie in large numbei'S around the female apei'ture. 

The female apparatus is no less remai'kable than that of the male. 
The vagina {va}) runs forwai-d for some little distance from the 
apertiu'e, then turns upwards. As it does so, it is twisted into 
a remarkable spiral coil, making some five complete turns. . It 
then runs backwards, narrows considerably, and soon receives the 
openings of the two uteri {ut.) on its ventral side. Beyond this 
point it is continued back as a narrow accessory vesicle {accves.) 
about as far as the level of the female aperture, when it tui-ns 
shai-ply ventralwards and oj^ens to the exterior hy the caitriwi. 

The only other Polyclads with which I am acquainted that 
possess a secondary female opening ai'e Trigonoporus of Lang [6] 
and Polyporus of von Plehn [11], but in both these cases the 
second opening is quite distinct from the primary one (the true 
antrum femininum). 

The curious spiral twisting of the vagina in the present species 
is, so far as I know, unparalleled in the oi-der. 

For the first part of its coui'se, %. e. whilst it is running forwards, 
the vagina is lined with elongated cokimnar ciliated cells, the nuclei 
of which lie near their bases. Outside this epithelium is a thin layer 
of circular muscle-fibres, and beyond these, in my sections, can be 
seen a great number of nuclei massed round the vagina, and 
probably belonging to gland-cells. Where the vagina turns 
dox'salwards and becomes spirally twisted it has narrowed slightly, 
but its epithelium i-etains the characters already mentioned. The 
muscle-sheath does not follow the individual folds of the spiral, but 
forms a continuous covei-ing for that part of the duct (PI. XXIII. 
fig. 6, m.s.). The rest of the terminal female ducts are precisely 
similar in character to the first part of the vagina, only narrower. 
There seem to be no special shell-glands present. 

The genus may be defined as an Acotylean genus in lohich behind 
the functional penis a small second penial organ occurs. The 
accessory vesicle of the female apparatiis opens to the exterior 
through the antrum femininum. Body pointed at either end. 
Marginal eye-spots present ; moibth subcentral. 

Family Latocestid^. 


Four specimens were collected by Mr. Evans. One of these is 
labelled " Chocolate-brown above, slightly lighter below." 

The largest specimen has a total length of about 30 mm. and a 
breadth of 5 mm. The mouth is 2*5 mm. from the hinder end of 
the body. 

This species is proportionately longer than the other member of 
the genus which I have had an opportunity of examining. Its 
most striking feature is the presence of a crowded row of eye- 
spots running completely round the margin of the body. The 




arrangement of the eyes at the anterior end of the body is shown 
in text-fig. 55 A and towards the hinder end in 55 B. 

The only other elongate form exhibiting this chai'acter with 
which I am acquainted is CestO'plana ? maldivensis niihi. This is 
ahnost certainly a Latocestus. 

Text-fig. 55. 

Eye-spots of Latocestvs aryus. 

I can detect no rhabdites in the epidermis. The parenchyma 
is very dense, and nervous tissue abundant on the ventral side 
just within the body-wall muscles. 

The terminal parts of the reproductive apparatus are almost 
exactly similar to those figured by von Plehn [7] for Latocestus 

314 MR. F, F. LAIDLAW ON THE [Apr. 21, 

As in that species, the testes as well as the ovaries lie on the 
dorsal side of the body. 

Thysaxozoon auropunctatum Coll. 

Thysanozoon aibropitnctatum Lang [6] ; von Stummer-Traiinf els 

One specimen, unfortunately immature and in a poor state of 
preservation, although it has retained its colour fairly well. 

PsEUDOCEROS BEDFORDi, sp. n. (Plate XXIII. fig. 9.) 

This strikingly handsome and large sjDecies was obtained by 
Messrs. Lanchester and Bedford in Singapore Hai-bovir. The 
single specimen, kindly given me by Mr. Lanchester, was unfortun- 
ately in fragments and fully one-half of the specimen missing, so 
that a satisfactoi-y examination was not possible. I believe, how- 
ever, that it was provided with a pair of penes. The colouring is 
very beautiful, and quite suificiently marked to render the identifi- 
cation of this species in the future a simple matter, thanks to the 
careful drawing for which I am indebted to Miss Dust. 

On the label accompanying the specimen is written " Singapore 
Harbour, from tide-marks to 10 faths." I venture to associate 
with this species the name of my lamented friend the late 
Mr. Bedford. 


Closely allied to Ps. cerebralis (Kelaart) and Ps. striatus 
(Kelaart), but smaller than either. It has the same i-egular 
convolutions at the margin of the body that are shown in 
Collingwood's figures, and the coloration is similar especially to 
Ps. cerehi'alis. 

Length, about 30 mm. 

Breadth ,, 25 ,, 

Mouth-opening, about 3 ,, from the anterior margin. 

Sucker about 8 ,, behind the mouth. 

The specimen is immature and the penis is unpaired. Colour — 
dorsal surface mottled dark brown and brownish white. The 
extreme margin is white, but the white rim is exceedingly thin ; 
just inside this is an equally fine black line. Ventral surface pale 
brownish white, becoming darker towards the margin, which is 
edged with black and white just as on the dorsal surface. 

PsEUDOCEROS ? RUBELLus, sp. n. (Plate XXIII. fig. 10.) 

A number of specimens of this very small species were collected. 
One of these was " found under a stone between tide-marks. It 
was magenta-red in colour. Xov. 1899." Though very small, 
these specimens appeal- foi- the most pai't to be mature. They 
bear a close resemblance to Pseudoceros kelaartii (Coll.) in colour, 
but the lattei- species is much larger and has the eye-spots arranged 
quite difi^erently. The arrangement in the present species is shown 


in fig. 10 of Plate XXIII. The tentacles also in Colliugwood's 
species ai-e nincli moi'e prominent. The penis is unpaired. The 
body is nearly circular and the sucker median. Total length 
about 5 mm., bi'eadth 4 mm. The ai-i-angement of the eye-spots 
is veiy diflfei-ent from that normally found in Pseiuloceros, but I 
cannot find any other charactei-s distinguishing it from that genus. 

At least two othei- species of this genus ai-e repi-esented in the 
collection. One of these is a very small individual of the 
F. superb us- gvouTp (perhaps P. hancockianus) ; whilst several 
specimens belong to a species coloui'ed exactly as Proceros con- 
cinnum Coll., viz., they are blue, with orange mai-gin and median 
stripe. They are, however, very much smaller, about 8 mm. to 
10 mm. in length, and of a different shape, not pointed at their 
hinder end, and have small, folded tentacles, whereas Collingwood's 
species has pointed tentacles. Proceros concinnum in fact is almost 
certainly a Euryleptid, whilst these specimens are undoubtedly 
members of the genus Pseudoceros. 

I prefer, howevei', not to name them at present. 

Family Diposthiid.e. 

AsTHEXOCEROS, gen. nov. 

AsTHENocEROS wooDWORTHi, sp. 11. (Plate XXIII. fig. 7.) 

Two specimens, without notes as to colour ; but, to judge from 
these spirit-specimens, the species is of a leddish-brown colour, 
darkest on the middle line. 

Length, about ... 17 mm. 

Breadth U „ 

Mouth-opening... 7 ,, from the anterior margin. 

Sucker 1*5 ,, behind mouth. 

The male and female apertures lie in the usual oi-dei- between 
the "mouth" and sucker. 

The pharynx is large and much folded, its opening being sub- 
median. This featui'e will serve at a glance to distinguish this 
species from any of the Pseudoceridfe. In describing the type of 
the family, Woodwoith [16] makes no statement as to the pharynx, 
but, to judge from the figure given, it is median oi' snbmedian as 
in the present species. 

Tlie bod}^ is flat and almost circular ; the anterioi- mai'gin is 
feebly folded, and thei'e is a small group of eye-spots on either 
side of the middle line, but there are no very definite tentacles. 
The prostate body lies in front of the jjenis. 

In this respect Asthenoceros diffeis fiom the type-genus Dipos- 
thus. There ai-e no biain-eyes. 

Only one of the two specimens was cut into sections, tx^ans- 
versely ; the other specimen appesirs, from an examination of it 
when cleared in oil of cloves, to be quite immature. The speci- 
men serially cut presents certain peculiar featuies, which I am at 

316 MR. F. F, LAIDLAW ON THE [Apr. 21, 

a loss to explain satisfactorily. In the first place, the testes and 
ovaries are very immature, and appai'ently only the terminal parts 
of the genital ducts are developed. On the othei- hand, there is, 
immediately over the penis, a large mass of mature spermatozoa 
lying in a chambei- the character of which cannot be determined 
from my specimens, since it appears in places to have indications of 
a proper lining-epithelium of its own, and again in places seems to 
be merely a gap in the parenchymatous tissue. There are very 
faint indications of a duct running from this chamber down in 
the dii-ection of the penis, but this duct cannot be traced far. 
Woodwoi-th [16] suggests in connection with the specimens of 
Diposthus described by him, that they were in a late stage of 
sexual activity, and that in consequence of this the sexual organs 
were in a reduced condition. Possibly in my specimens of 
Asthenoceros a similar state of afikirs occurs, but, as I have 
already stated, testes and ovaries in a very immature sta,te are 
present. There is also a possibility that the ripe spermatozoa 
may be derived from another individual by hypodermic injection ; 
but I do not think this is the case — firstly, because the penis is 
not armed with a stylet ; and, secondly, because the spermatozoa 
lie over the penis, and because, as already stated, there are traces 
of a duct running towards the penis from the chamber in which 
they lie. 

The terminal parts of the male apparatus histologically resemble 
those of BipostMts. The prostate and penis are separated into 
two distinct organs, " both of which are doubtless intromittent," 
both opening by a single gonopore. The prostate lies immediately 
in front, the penis directly over the aperture. It is rather 
feebly supplied with muscle-fibres, and the secretory cells lie in 
the middle of the organ, but there is no lumen apparent 
(cf. DipostMis corcdlicola). 

The penis is much more muscular, and on the outer side has a 
very definite series of cii'cular and longitudinal fibres. Nuclei in 
it too are much more niunei-ous (PI. XXIII. fig. 7). 

In connection with the female organs there are traces of two 
pairs of viterine vesicles. The uteri are very small and difficult to 
distinguish from the surrounding tissue, and the ducts running 
to them are merely solid rods of cells of an embryonic appearance. 
The uteri open into the hinder end of the vagina. The antrum 
is deep, and widens at its upper end where it receives the secretion 
of the shell-glands. 

The body epithelium is very densely crowded with small 
I'habdites, and on the dorsal side especially with pseudoi-habdites. 
Scattered through the parenchyma, more particularly in the region 
of the sucker, are numbers of small rounded darkly-staining cells 
the nature of which is doubtful. The ventral surface of the body 
projects immediately behind the pharynx into a prominent median 
ridge which carries the gonopores and the sucker ; behind the 
sucker the ridge disappears. 

I have given as complete an account as possible of the cha- 


racters of the genital organs, but, owing to the comlition of tlie 
specimen, it is obviously fai- from being satisfactory, and in order 
to deal fully with this interesting species more material is 

Prosthiostomum pallidum, sp. n. 

One specimen, from tlie sea-shore, Dec. 1899. 

Total length, about 20 mm. 

Breadth, about 4 ,, 

Anterior margin I'ounded. Arinxngement of eye-spots shown 
in text-fig. 5G. 

Text-fio-. 56. 

Ej'e-spots of Prosthiostomum pallidum. 
e, margiual ; br.e, bi-aiii ej'e-spots. 

Closely allied to P. siphancuhis of the Meditei-i-anean by the 
ari-angement of the eye-spots, and agreeing with it apparently in 
being, so far as the spirit-specimen shows, of an uninterrupted 
dull grey-bro-wTL colour, it differs sufficiently in that the two rows 
of brain-eyes diverge continuously from one another from before 
backwai-ds, whilst in the Mediterranean species these two rows 
converge at their middle. 

It is also readily distinguishable, I think, from the latter species 
by its smaller size. The single specimen obtained by Mr. Evans is, 
to judge from the state of the sexual apparatus, fully mature. 
P. siphitncidus, moreovei', has the genital sexual organs relatively 
much smaller, to judge from Lang's figui'es (see Lang, ' Poly- 
cladida,' pi. 5. fig. 3). 

P. pccUidicm Ls certainly veiy distinct from any species from 
the Indian Ocean that I have had the opportunity of studying, 
and also, I believe, from the species found in the Pacific. 

I believe that Leptoplana aii,rantiaca of Oollingwood is really a 
Pi-ostkiostomum. It has the shape and 2)roportions of a member 
of that genus, whilst its eye-spots have the characteristic Pros- 
thiostomuiii arrangement. It may, of course, be identical with the 


species described above, but, judging from Colling-wood's figui-e, 
in the arrangement of the eye-spots it is sufficiently distinct. 

Liter atttre. 

1. Bergendal. "Kongl. Fysiogr. Sallskapet," Lund Handlingar, 

Ny Foljd, ] 892-93, Bd. 4. 

2. CoLLiNGWooD. Trans. Linn. Soc. Lond. ser. 2, Zool. i. 1875, 

p. 83, pis. 17-19. 

3. Graff, von. Zeitschr. wiss. Zool. Iv. 1893, p. 189. 

4. Laidlaw. Fauna and Flora of the Maldives and Laccadives, 

i. p. 282, pis. xiv., xv. 

5. Mem. & Proc. Lit. &, Phil. Soc. Manchester, xlvii. p. 1 


6. Lang. Naples Monogr. xi., 1884. 

7. Plehn, von. Jena. Zeitschr. xxx. p. 137, t. viii. & xiii. 

8. Ergebn. Plankt.-Exp., Heft i., 1896. 

9. Semon's Zoologische Forschungsreiseil, Bd. v. pp. 329- 

334, t. xxiii. 

10. Abh. Senckenb. Gesell. xxiv. ii. pp. 145-146 (1897). 

11. Jena. Zeitschr. xxxi. p. 90, t. v. (1898). 

12. Zool. Jahrb. Syst. xii. pp. 448-452 (1899). 

13. Stummer-Traunpels, von. Zeitschr. wiss. Zool. Ix. p. 688, 

t. xxxv.-xxxviii. 

14. Verrill. Trans. Connect. Acad. viii. p. 459, pi. xliv. 

15. ,, „ ix. p. 523. 

16. Woodworth. Bull. Mus. Oomp. Anat. xxxii. 4, pp. 63-67, 

pi. i. (1897). 


Fig. 1. Spine from the lumen of the penis of Flanocera sp., p. 302. 

2. Section passing through the penis and prostate organ of NotoplMm evansii, 

p. 302. 

3. Transverse section through the penis oi Leptojplana malayana, p. 306. 

4. Transverse section througli the penis and prostate gland (?) of Bergeiidalia 

anomala, p. 310. 

5. Section through the rudimentary organ of penis-like structure of the same 

species, p. 311. 

6. Section through the spiral part of the vagina of the same, p. 312. 

7. Section across the antrum masculinum of Asthenoceros tooodivorthi, p. 315. 

These organs are directed obliquely forward, and consequently both appear in 
some of the sections. The granular appearance of the prostate in this region 
is rather exaggerated in the figure, but it becomes more marked towards its 

8. Sergendalia anomala, p. 306. 

9. Pseudoceros bedfordi, fragment of anterior part of body, p. 314. 
10. Pseudoceros ? rubellus, p. 314. 

Explanation of Lettering of Plate and Text figures. 

a.m., antrum masculinum. 
a.m.l., muscular walls of antrum mascu- 
d.e., ductus ejaculatorius., prostate duct. 
ep., epidermis., muscular walls of prostate. | t., tentacles. 

pr.c, chamber of the prostate. 
pro., prostate. 
ppr., prostate-like organ. 

s., septum between chamber of 

St., stylet of penis. 

m.v., muscular sheath of vagina. 

p., penis. 
pp., vestigial penis ! 
pr., prostatic cells. 

va., lumen of vagina. 
v.d., vasa deferentia. 
V.S., vesicula seminalis. 


4. On the Phyloo-enetic Cnnso of tlic Transposition oF tlie 
Testes in Mammal la : with Remarks on the Evolution 
o£ the Diaphraom and the Metanephrie Kidney. By 
W. Woodland, University College, London.' 

[Received March 3, 1903.] 
(Text- figure 57.) 

Tlie sulijtK;t of th(! descent of the testes in ]\riimiu:di!i, notwitli- 
standing its biononiic intori'st, has liitherto rt'ceived but littk^ atten- 
tion from the phylogenetic standpoint ; indeeil, so far as 1 know, 
its intei'protation has not hitherto been attempted. To exhibit 
the general nature of the problem and the apparent dilHculty 
attending its solution, tlie following paragraph from Mr. Spencer's 
'Principles of Biology' (vol. i. p. 573) may be quoted : — "But 
now let it be confessed that though all phenomena of organic 
evolution nuist fall within the lines above indicated, there remain 
many unsolved problems. Take as an instance the descent of the 
testes in the Mammalia. Neither direct nor indirect equilibration 
accounts for this. We cannot consider it an adaptive cliange, 
since thei'e seems no way in which the production of sperm-cells, 
internally carried on in a bird, is made external Vjy adjustment to 
the changed requii-ements of mammalian life. ISTor can we ascribe 
it to survival of the fittest ; for it is incredible that any mammal 
was ever advantaged in the struggle for life by this changed 
position of these organs. Contrariwise, the removal of them 
from a place of safety to a place of danger would seem to be 
negatived by natural selection. Nor can we regaixl the tiuns- 
position as a concomitant of re-equilibration ; since it can hardly 
be due to some change in the general physiological balance." I 
agree with Mr. Spencer that the descent of the testes can neither 
have been a change adaptive in nature, noi- a result of the 
operation of natural selection ; on the other hand, I must dis- 
agree with the statement that the phenomenon cannot be 
regarded " as a concomitant of re-equilibivition," since, as will be 
shown, I attribute it to the direct action of the conditions of life 
found in the Mammalia. 

To ensure due appreciation of the significance of the following 
statements, it will be as well to here indicate the nature of the 
theory about to be advanced. Of possible causes efi"ecting the 
transposition of the testes, there are two categories : (a) causes 
which as regards their eftects on the oi'ganism mediately 
or immediately impart advantage, so supplying the requisite 
material for natural selection ; and (6) causes which in their 
eflfects on organisation bear no appreciable relation to either 
advantage or disadvantage. Since those of the former categoiy 
are inconceivable in this connection, we are led to conclude 

1 Communicated by Prof. E. A. Minchin, F.Z.S. 
Pkoc. Zool. Soc— 1903, Vol. I. No. XXI. 21 

320 MR. W. WOODLAND ON THE [Apr. 21, 

that the transposition is an inevitable concomitant of some other 
constant feature of the animal's existence, thus not having arisen 
in relation to ulterior ends. Again, since the testes consist of 
ordinary matter possessing inertia and mass, their translation 
(involving rupture or distension of the mesorchium) implies 
mechanical force. From both of the foregoing considerations, 
and from the fact that no other efficient cause is imaginable, 
it is probable that the displacement results from the reaction 
en the part of the testes to the incident forces, which, it can 
be shown, are generated by bodily activity under mammalian 
conditions of life. The theory here advocated is to the effect 
that the descent of the testes in the Mammalia has been pro- 
duced by the action of mechanical strains causing rupture of the 
mesorchial attachments, such strains being due to the inertia of 
the organs reacting to the impulsiveness involved in the activity 
of the animals composing the group. 

Mechanical Aspects of Organisation. 

Before entering upon a discussion as to the production of the 
forces above mentioned and the manner in which they have 
acted, it is well briefly to outline the entire argument from the 
mechanical standpoint. In the first place, accelerations imparted 
to the body as a whole generate strains (or stresses) in connection 
with the attachment or other means of support of every com- 
ponent viscus, and the degree of such accelerations (and therefore 
of the strains and stresses) is obviously dependent upon the 
reaction which occurs between the animal's body and the medium 
in or substratum on which it is supported, being directly pro- 
portional to the product of the powers of resistance possessed by 
the substances constituting the same. Secondly, it must be 
pointed out that the accelerations to which we refer, and to 
which alone we attribute any importance, are those involved in 
the impulses communicated to the body during the actual con- 
tinuance of locomotion, and which inevitably result fiom the 
mode of action of the propelling agency, whatever its nature. 
The ordinary non-impulsive accelerations involved in the starting 
of an animal into motion from a state of rest, or its converse, do 
not concern us. In considering, however, the arboreal habits of 
e.g. Primates, the case is different, accelerations involved in 
motion from rest here being decidedly impulsive in nature. Now, 
among the Vertebrata, it is evident that fishes, aquatic amphibia, 
and birds severally exist in media which, owing to their mobility, 
negative the occurrence of accelerations of high degree, i. e. 
impulsiveness. Hence, as regards these accelerations, we have 
to consider only terrestrial animals, which come into contact with 
a substratum possessing sufiicient power of resistance to afford a 
reaction of mailced intensity. Of these terrestrial animals the 
mammaliaii group is at once distinguished from terrestrial 
Amphibia and Reptilia, both by the high degree of activity which 


its members exhibit, and by the moi'c pci'fect adfiptatiou of tlic 
body (relation of limbs to trunk, &c.) to the locomotion in which 
the superior activity is mainly manifested. Consequently, in 
mammals, the reactions which occur between the organism and 
the substratum vastly exceed in magnitude those occurring in 
lower gi'oups, and we have in this distinction a factor which in 
considering bodily confoiniation is worthy of all attention. 
Since in mammals tlieorgnnisMtion is subject to such considerahle 
forces, it follows that if stal)ility of position Ijc required in con- 
nection with organs possessing jipjjreciaLle mass, i. e. if these 
organs are to adjust themseh es to those transposing inlluences 
which arise out of inq^idsiveness, fixative structures must bo 
developed in response to the demand. For, as remarked above, 
a sudilen accelei-ation imparted to the body is necessarily trans- 
mitteil to an attached organ through its attncliment, which, if the 
organ be of consideiublc mass, is thereby su1)jected to an intense 
strain, possibly ending in disruption. Stability of position of an 
organ may be necessitated either on mechanical or physiological 
grounds : thus, apart from considerations of function, viscera of 
large mass (stomach, liver, intestine) must conform, as regards 
situation in the body-cavity, to the principle that, cceteris paribus, 
the more anterior the centre of gravity of the body the greater 
the facility of progression ^ ; and again, from the functional 
standpoint, the kidneys must in active animals be maintained 
anteriorly in order to ensure j)roximity to a vigorous blood-suj)ply. 
It will be evident from this, that if the position of an organ be of 
no economic moment — if thei'e be no need of localisation either on 
account of mass or volume for preservation of bodily equilibi'ium, 
or on account of nature of function for j)hysiological etticiency, — 
retentive structures will not tend to be formed, and the organ, if 
of sufficient mass, will lespond by transposition to any consideralile 
force incident upon it — a trait which in the course of generations 
will be inherited. 

I shall no^v proceed to examine more in detail the production of 
the forces hitherto assumed to be capable of eflecting the trans- 
position of the testes— to enquire whether the intensity and 
direction of these forces are such as are congruous with the 
hypothesis advocated. 

Mammalian Locomotion. 

In view of the importance that is here attached to the character 
of mammalian locomotion, it is needful to discuss the sidjjcct in 
detail. Every unsupported mass descends to the earth with a 
given acceleration, and the magnitude of tlie force required to 
re-elevate the body to its initial position is directly propoi-tional 

1 Well shown by measurements of the position of the centre of gravity in Pisces, 
and by the abdominal contours of active cursorial mammals as compared with those 
of species more slothful in habit (compare e.^. a greyhound or race-horse with a 
St. Bernard or cart-horse, or, more generally, lierbivores witli ciirnivores, though 
the dlflerent capacities of the digestive organs have hcix- to be ooiisid;roil). 


322 MR. W. WOODLAND ox THE [Apr. 21, 

to the height tliroiigh which upheaval is effected, and inversely 
proportional to the period of time occupied by upheavaL This 
statement in its application to terrestrial locomotion provides 
the clue for the solution of the present problem. With the 
exception of aquatic organisms, which exist in a medium so 
similar in density to themselves that the influence of gravity is 
not felt, all animals which fly, crawl, leap, or run are subject to 
this primary condition of self-support imposed by the earth's 
attraction. In Aves the body is not entirely unsupported, the 
resistance of the air to the large expanse of body yielding con- 
siderable aid in this respect, and the small amount of self- 
sustainment needed can be provided by the minor elevations 
imparted by individual strokes of the wing. The reactions 
between the wing and the air are of necessity small, owing to the 
mobility of the latter substance, and hence, in spite of the great 
activity of birds, no great accelerations are imparted to the body 
— the locomotion is not impulsive. The conditions affecting 
terrestrial locomotion are exceedingly unlike. Here, as before 
remarked, the powers of resistance possessed by the substances of 
oi-ganism and substratum both being of high degree, reaction 
between the two is correspondingly great. In Reptilia and 
(excepting the saltatory Anura referred to below) terrestrial 
Amphibia, however, the forces involved in locomotion are not 
conspicuous for their intensity, owing both to the fact that the 
period of upheaval is prolonged (no sudden impact occurring 
between the limbs and the earth) and the small amount of eleva- 
tion effected by the action of the limbs, these two features 
resulting from the conformation of the body and the general 
inactivity of the animal. In Mammalia, on the other hand, the 
period occupied by contact of the limbs with the earth is 
extremely biief and the height of elevation considerable ; and 
hence, though the angulation of the limbs tends to diminish 
concussion, the organisation is subject to the intense strains and 
stresses resulting from the enormous forces generated during 
locomotion. Consider the gallop of a typical Ungulate or 
Carniv'ore. The elongated trunk possessing two pairs of limbs, 
each pair being in an opposite phase of motion compared with 
the other, it follows that the two halves of the trunk will alter- 
nately be upheaved through a considei-able distance on contact 
with the earth being made by their respective paii-s of limbs, and 
depressed in the interval which exists between successive contacts. 
That is, the pendulous swing of each pair of extremities is accom- 
panied by an elevatoiy impulse at the centi'e of the ai-c each 
describes, and these impulses effect the upheavals of the respective 
halves of the body through the distances wdiich they have fallen 
in the time required for each pair of limbs to describe twice the 
length of its ajDpropriate path. Oonsideiing the mode of action 
of either pair of limbs, and beginning at the horizontal stage of 
the trunk's position when the half of the body is being depressed 
under the influence of gravit}^, this continues to descend until the 

1903.] TKAxsrosixiox of mammaliax testks. 323 

limbs make contact with tlie earth, when, as alreail}' descri^jed, 
the entire portion of the trunk is suddenly u])heaved with con- 
siderable force. If the horizontal velocity could be reckoned as 
constant, the only accelerations which the trunk would acquire 
would be in a vertical line ; but since the hoi-izontal velocity is a 
quantity which, owing to the i-esistance of the air, is continxially 
decreasing, the upward acceleration suddenly imparted to the 
trunk at each contact of the limbs with the earth has to be com- 
pounded with a sudden forward accelei-ation in order to maintain 
the pace, and lience the resultant acceleration which the half of 
the body acquires at eveiy upheaval is in an apu-ard i\\\(\ for tear d 
direction. It is most essential to I'ecognise this fact that the body 
as a whole has periodically imparted to it large accelei'ations in 
these two directions. As is obvious, the upwai'd and horizontal 
velocities due to these accelerations become respectively nei^tralised 
by gravity and atmospheric resistance at the middle and enrl of 
the interval which exists between successive contacts of the limbs 
with the earth. From the instant when the maximum elevation 
of the half of the trunk is attained, i.e. when the upward velocity 
is completely neutralised, until contact again occurs, the body 
continues to acquire a downward acceleiution due to gravity. 
But on contact taking place, the lai-ge downward velocity due to 
this acceleration is suddenly destroyed, and as suddenly exchanged 
for an equal upward velocity, in the biief instant of time occupied 
by the impact of limbs and earth. Whence it is evident that the 
degr-ee of upwaixl accelei'ation imparted to the half of the body 
during the sudden loss and gain of downward and upward velocities 
must be considerable. It isnoteworthy that whereas each /br^t•<:«•f^ 
acceleration, due to either pair of limbs, is imparted to the body 
as a whole, each upioard, acceleration is solely imparted to that 
poition of the trunk to which the pair of limbs causing the 
elevation belongs ; and hence the attachments of those oi-gans 
situated in the median portion of the trunk are evidently not so 
subject to downward strains as in the case of organs placed more 
tei-minally. Tlius it is important to notice, with regai'd to what 
follows, that the primitive pre-mammalian portion of the testes is 
decidedly posterior (see diagram, p. 334), and that in consequence 
the motion of the hind portion of the body is the factor of most 
importance in the consideration of the present problem. 

From the foregoing, it is sutHcienth'- manifest that at each 
elevation of the body in maunuals a considerable force is imparted 
to the organisation as a whole (and by necessary implication to its 
constituent parts) in an upward and foi'waixl direction — as was to 
be proved. If the constituent pai'ts of the body be considered, it 
Avill be evident that such a force woi'ks differential effects among 
them. For these organs, lai'gely diflering among themselves in 
regard to mass, and the same force being incident on all, it follows 
that the individual reactions are different, and hence there exists 
a tendency to segregation of those organ;; of greater mass from 
those of less. Moreover, oi-gans differ inter se in their relation 

324 MR. W. WOODLAND ON THE [Apr. 21, 

to the rest of the body. If an organ be imbedded or otherv/ise 
firmly affixed to the main bulk, it Avill on account of its large area 
of attachment and close apposition readily share both the eleva- 
tions and depressions incurred in locomotion — the strains and 
stresses are distributed in space and time, and hence their inten- 
sity is lesseiied ; if, on the other hand, a massive organ be merely 
suspended fi'om the main mass of the body, such a condition 
subjects the attachment to the influence of severe strains periodi- 
cally recurring, the direction of Avhich approaches that of the 
resultant acceleration of the trunk during its impulsive elevation. 
The great severity of these strains on the attachment of an organ 
thus disposed will be more fully realised if it be remembered 
that, on the sudden elevation of the body, the organ possesses the 
downward velocity due to the previous depression — a momentum 
which in being destroyed throw^s additional strain on the means 
of support. As will be seen, the preceding remarks apply in an 
eminent degree to the testes. 

The Genitalia of Mammalia. 

The reproductive organs of the Mammalia are relatively small 
bodies of great density and, in the case of the males, compactness, 
and they primitively occupy a position similar to that found in 
the lower Vertebrata. It is, however, characteristic of the majority 
of the Mammalia that in the course of development the testes 
forsake their primitive lodgment, and migrate posteriorly and 
ventrally to the terminal periphery, where they protrude at the 
surface of the body- wall. This protrusion constitutes the scrotum, 
in the wide sense of the term, which may vary in character from 
that of a pair of small slightly-elevated areas to that of a capacious 
oval pedunculated sac. Apparently in man alone the section of 
the ccelom contained within the scrotum becomes completely 
separated oft' from the main cavity ; in other mammals, com- 
nivmication is retained by means of the inguinal canal, which, 
however, is usually narrow, so negativing return of the testes to 
the main body-cavity. This feature in the case of active 
mammals possessing large testes is very important, since if the 
unattached organs were permitted to return to the main cavity, 
the forces to which they would be exposed during locomotion 
Avould doubtless be exceedingly detrimental — a malinfluence which 
both the narrowing of the inguinal canal and the possible tractive 
function of the gubernaculum during development safeguard 
against. In fact, contrary to the usual supposition, the internal 
and not the external situation of the testes is the source of 
danger. It will also be observed that in small animals which do 
not possess nai'row inguinal canals, there is no such dire necessity 
for the resti'iction of the testes to the scrotal cavity, both on 
account of the smaller size of the organs and the usually less 
intense forces to which the testes are subjected. Thus, para- 
doxical as it may seem, the increased protrusion of the testes 


beyon?l the surfcxce of the bo;ly-wall, besiles being a necessary 
result of more massive oi-gans and the incidence of more powerful 
forces, is in itself a further means of preservation. In physical 
character, the testes, as already mentioned, are definite bodies of 
concentrated form, suspended dorsally in the body-cavity by the 
thin mesorchial membrane. Also investigation shows that, with 
the exception of the heart, the testes are the densest oi-gans in 
the vertebi'ate body, and hence, as they possess appreciable volume, 
their mass is also considei'able. Moreover, the testes (and ovaiies) 
are distinctly separated from the remainder of the body, this 
separation (lending additional facility to transposition) arising 
out of the fact that the gonads ai-e fi-om their veiy nature indi- 
vidualised, reproduction essentially consisting of the separating 
oil" of a portion of the oi'ganism ; and hence the body prinuuily 
serves as a mere carrier of these organs, which, unlike the 
kidneys e. g., bear no relation to the economy of the animal. 
From this it follows that the transposition of the testes is of no 
concern to the rest of the organism, since the process can cause 
no derangement of function in other poi'tions of the body ; and 
hence the testes, differing from all other organs in this respect, 
do not, under oi'dinary conditions, reqidre to be retained in their 
])rimitive position by the special development of fixative structures. 
Thus, as regai'ds their definiteness and concentration of form, 
their means of suspension, their superior density, and their 
structural and physiological separateness fi'om the i-est of the 
organism, the testes fully coaiform to the above-specified con- 
ditions favourable to transjDOsition. The transposition of the 
testes occuning under conditions which permit the descent of 
these organs alone (and the testes alone have descended), and a 
cause capable of effecting this transposition solely existing in the 
Mammalia (in which group descent has alone occurred), it is 
probable that the latter phenomenon is the cause of the former. I 
hold that, in the majority of the Mammalia, the testis attachment 
has throughout tlie histoiy of the race been constantly subjected 
to severe strains consequent on the character nnd conditions of 
mammalian locomotion, and that on account of the resulting 
disruption or distension of the mesorchium, the testis has migrated 
in a postero-ventral line {i. e. in an opposite direction to the 
forward and upward accelerations imparted), coming to lie at the 
terminal periphery of the body- wall and forming the scrotal pro- 
trusion. Just as when a man runs, a weiglit in his coat-pocket 
will periodically " drag " and ultimately wear a hole in the lining 
by constant distension, so the testis of mammals has responded to 
like forces resulting in " descent." 

I now proceed to consider the genital orgnns of the Mammalia 
as a Avhole, the general conformation of which amply confirms 
the foregoing conclusion, tending to show that the testes have, 
in evei-y case, reacted in a degree proportional to the forces 
concerned. Indeed, the general correspondence between situation 
of the testes and grade of impulsiveness displayed in the various 

326 MR. W. WOOBLAND ON THE [^^V^'- 21, 

oi'dei's and families of the Mammalia affords such conclusive 
evidence as to the causal relation subsisting between the two 
that a systematic review of this evidence is well called for. That 
the coi'i'espondence is not absolute, however — that there exist 
instances of the concurrence of scrotal testes with sluggish habits 
— is without doubt mainly, if not wholly, to be explained by 
phylogenetic considerations. For it must be remembered that 
once the descent of the testes is inherited, the trait is a constant 
one (unless acquired sluggishness is able in time to produce efTect, 
which is doubtful), and remains so in successive generations 
whatever varied habits may be assumed, unless the bionomic 
aspect so changes that natural selection cancels the " plus varia- 
tions," so affording an ascendancy to reversionaiy factors. That 
such has been the case in several instances will be illustrated 

In the Monotremes the activity (impulsiveness) is of a veiy 
low degree. Ornithoi^hynchus is " aquatic in its habits, passing 
most of its time in the water or close to the margin of lakes and 
streams." The " body is rather long, compact, and almost every- 
whei"e of the same thickness. It rests on short, massive legs 

so shoi't that the animal in walking or running actually 

drags its body along the ground." Echidna is " usually found in 

I'ocky districts, and more especially in the mountains and 

is mainly of nocturnal habits." It is described as " indolent." 
The testes in both instances are " abdominal in position through- 
out life," and afford the only example in mammals of a disposition 
of these organs anterior to the kidneys. The low status of these 
animals is well known. 

In the Marsupials " the testes are always contained in a scrotum, 
which is susjoended by a narrow pedicle to the abdomen in front 
of the penis." If it is permissible to assume that the ancestor of 
the marsupials was kangaroo-like, the pre-penial position of the 
testes may perhaps be attributed to the peculiar mode of loco- 
motion characterising this animal. Foi- the rapid locomotion of a 
Kangaroo consists of a series of leaps, and such leaps would involve, 
as in ordinary mammals, a series of antero- dorsal tensions on the 
suspensory membranes of the testes ; but since the bodily depres- 
sions ai-e mai'ked by a more prolonged termination, as compared 
with those involved in the ordinary gallop, the testes would in 
addition tend to be thrown' forward, and these two factors in 
conjunction have possibly led to the peculiar position of the testes 
found in marsupials. On no othei- theory than that of descent 
from an ancestor characterised by jjre-penial testes can the peculiar 
genital conformation of the variously habited marsupials be corre- 
lated Avith special bionomic conditions ; and hence, on this ground 
alone, it is possible, as implied above, that the ancestor of the 
Marsupialia was of the type of the Macropodidse, the characteristic 
Saltatory progression originating the pre-penial situation of the 
scrotum in tlie manner indicated. 

The Edentata repi'esent anothei- ancient gi'oup. Their activity 


in genei'.'il is siimll. Tlie Brarlypodidse are arboreal in habit, 
and are chai'acterised by " habitual sliiggishne,ss," being '• most 
inefficient walkers," and in climbing, never leaping from bough 
to bough (a form of activity invohing more impulsiveness than 
even the gallop, but probably not more than that involved in 
saltation). Their testes ai'e placed close to each other, lying on 
the rectum between it and the bladder; i. e. are retained in the 
abdomen. The Myrmecophagidaj are also not noted for activity, 
and their testes are disposed as in the Sloths. The Dasypodidie 
ai'e similarly inactive, being " harmless and inoffensive," endeax'our- 
ing to escape by rapid burrowing. However, it is stated that 
" they can run with great rapidity." The testes are abdominal, 
lying "above the brim of the pelvis." The Manidje are terrestrial 
and burrowing in habit, but some members of the gToup can 
climb trees. They are not very active. The testes lie in the 
inguinal canal. The OrycteropodicUe (Aard-vai'ks or African 
Ant-eatei's) are terrestrial and fossorial. Their testes ai'e 
" inguinal, but they appear to descend, at all events temporarily, 
into a scrotum." Their phylogenetic position is uncertain. In 
the order Sirenia the testes, needless to say, are abdominal. 
These animals are " slow and inactive in their movements, mild, 
inottensive," browsing at the bottom of watei'. 

The piscine locomotion of the Oetacea sutHciently accounts for 
the abdominal position of their testes, which organs are placed in 
the pi'oximity of the kidneys. Even assuming the tei-restrial 
ancestor of the Cetacea to have been charactei-ised by the exter- 
nality of the testes (which is improbable considering their existing 
localisation), such Avould inevitably have assumed an internal 
position consequent on the serious risk of injury involved under 
Cetacean conditions. 

The Ilodents are comparatively small animals. In habit they 
ai-e mostly teri'estrial , but some are arboreal and some natatorial. 
" The testes in the rutting-season form projections in the groins, 
but (except in tlie Duplicidentata) do not completely leave the 
cavity of the abdomen," i. e. the scrotal elevations are not 
well-defined. The Duplicidentata compiise the Hares, Rabbits, 
and tlie Picas or Tailless Hares, all extremely active animals, 
the latter being described as " agile " and as living in crevices 
among rocks. The Simplicidentata comprise the remaining 
Rodents, the principal families of which are the following : — 
The Sciui'ida3 " vary between the two extremes presented by our 
ordinary squirrels, the agile climbers, and the sluggish, clumsy 
marmots, which live almost entirely underground." Assuming the 
statement to be correct that the condition of the testes is similar in 
each of these two divisions (which is doubtful), it is evident that 
the latter are specialised forms descended from active ancestors. 
The locomotion of the Dipodidte resembles that of the marsupial 
Maci'opodidjie and insectivoi'an j\lacroscelid;e. " The whole struc- 
ture is adapted for jumping, and we iind resemblances in their 
btructui'e on the one hand to the kangaroos, and on tlie otlier 

328 MR. W. WOODLAND ON XUE [Apr. 2], 

hand to the jumping- shrews among the Insectivora." These 
animals, unlike the Macropodicla3, move their hind limbs alter- 
nately in the pi-ocess of walking. Unfortunately the writer has 
been unable to discover detailed information with regard to the 
testes in these groups. Other families are the clumsy, thick-set 
Hystricidae, the very active Castoridse (Beavers), Myoxidee (Dor- 
mice), and Murid^e. During the rutting- season the testes of 
Rodents migrate more posteriorly than at normal times ; but this 
merely indicates a slight displacement probably due to the periodic 
enlargement of the organ, the cremaster muscle eifecting return 
by decreasing the capacity of the scrotal emergence. This can be 
well seen in the Muridce, where the slightest impact will cause a 
dislodgment of the testis. 

The order Insectivora comprises "small animals of very 

low type ...... belonging to the oldest mammalian stocks 

which are generally terrestrial, although rarely of arboreal or aquatic 
habits the greater number are cursorial." '• In the sub- 
family Centetince, and Chrysocldoris, the testes lie immediately 
behind the kidneys, but in others more or less within the pelvis ; 
during the rutting- season they become greatly enlarged, forming 
protrusions in the inguinal region." The various families of this 
group afford good illustr-ations of the correlation of impulbiveness 
with degree of testis transposition. As stated above, in the 
Centetince or " Crawlers " — the appellation denoting their mode of 
locomotion — and Chi-ysocMoris, which is inactive and fossorial, 
the testes lie just posterior to the kidneys, i. e. are abdominal. 
In the companion subfamily, the Oryzorictince, the two genera 
are represented by small animals — Macrogale being mouse-like 
with a long tail, and Oryzorictes mole-like in form. The testes 
are situated near the urethra. In the Erinaceinm or Hedgehogs 
the testes are situated on the " underside of the inguinal canal." 
Their " movements are sluggish, their steps almost tottering, their 
gait clumsy." On the other hand, in the active Soricidas (Shrews) 
and saltatorial Macroscelidte (Jumping-Shrews), the testes project 
at the periphery of the perinteum, and in the Solenodontidte, the 
feet of which are "formed for running," the "testes are received 
into perinatal pouches." 

In the order Ohiroptera, the members of which are so highly 
specialised for flight, there is " no scrotum, and the testes are 
either abdominal or inguinal." " "We find in the low organization 
of their brain a proof of their inferior status " — a fact otherwise 
implied by the absence of the scrotum, unless a secondary reten- 
tion has occurred owing to the danger of partial externality 
involved in the conditions of flight and position of the hind limbs. 

The order Ungulata is subdivided into the Ungulata vera and 
the Subungulata. In the former suborder, needless to say, the 
testes of the large majority reside in a well-defined scrotum, cor- 
responding to the eminent impulsiveness of the animals. In the 
Nasicornia (Rhinoceroses), however, the testes are inguinal, the 
tunica vaginalis communicating freely with the body- cavity. As 


is well known, they five " luigo, heavy, clumsy creatures, with 
bent legs so short that the belly seen)S almost to drag on the 
gronnrl." The Subungulata comprise the Hyracoidea and the 
Proboscidea. The ancestry of the mendiers of both these sub- 
orders is as yet undecided, their status, decidedly low in the scale, 
being obscure. The Hyracoidea are animals of about the size of a 
rabbit or somewhat larger, and " resemble small maiinots." They 
possess a " short, fat body " with "weak and short feet." "In 
most species there is a complete adaptation to a life among the 
rocks," by possession of curious clinging habits resembling those of 
the Geckos. They are " agile in their sports, but lather lazy where 
food is abundant." Owen states that " the testes are abdominal, 
below or beyond the kidneys." The low status of the Proboscidea 
is shown by several anatomical traits, such e. g. as the possession of 
two venie cava? and the structure of the limbs. These animals 
have been well described as " peaceable colossi." Their gait is 
" pretty slow, though the Colossus can van very fast when once 
in full career, but this pace never lasts very long and is always 
maintained in a straight line." Owing to their huge size 
Elephants never gallop. The most active niendiers are the 
" I'ejected males" — such extiu activity thus being neutral so far 
as the inheritance of any tendency to testis descent is concerned. 
Elephants are vegetable feeders and ai'e " gregarious, generally 
inoliensive and even timid, fond of shade and solitude and the 
neighbourhood of water." The testes are permanently abdominal 
—a fact explicable by habits, though phylogeny alone can aflbrd 
a complete solution. Here, as possibly also in the case of the 
Nasicornia, it may be pointed out that the huge size of these 
animals itself implies inactive ancestors, for, according to one of 
the conditions of gTOwth enumerated by Spencer, great activity is 
antagonistic to increase by bulk, and the occuirence of the latter 
negatives the past existence of the foimer. 

The order Cai-nivor.a is subdivided into the Fissipedia and the 
Pinnipedia. The mend^ei's of the former group are all exceedingly 
active animals of large size, and, together A\ith the Ungulata, include 
the swiftest of terrestrial animals. Their testes, needless to say, 
reside in a well-defined sciotum. The Pinnipedia aftbrd an inter- 
esting illustration of the secondaiy operation of natural selection. 
In the Otariida; the " hind feet are turned forwai'ds under the body, 
and aid in supporting and moving the trunk as in ordinary 

mammals They spend more time on shore, and range 

inland to a greater distance than the true seals." In the Otariidje 
the testes are " suspended in a distinct external scrotum" '. On 
the other hand, in the Phocida^, "the hind limbs are directed so 
far backAvards that they continue the hoiizontal direction of the 

vertebral column They move on land only with difficulty 

by fixing themselves with their flippers in front and pulling up 
their hinder parts, then drawing their bodies up into a curve and 

' Owen states that the bcrotuni is uot distinct. 

330 MR. W. AYOODLAND ON THE [Apr. 21, 

throwing their front parts again forwards. They drag their belly 
along the earth, show little suppleness, and soon become tired." 
There is no scrotum, the testes being abdominal and " imbedded 
in areolar tissue." Thus, as befoi'e maintained, the embryonic 
ti'ansposition of the testes tends to be checked if external conditions 
become so changed as to render complete descent of these organs 
disadvantageous to the animal, such being obviously the case in 
the Phocidae, which have become further modified for an aquatic 
existence than the Otariidfe. It may be objected that ordinary 
mammals are subject, though in a less degree, to a like drawback 
— that their testes are also exposed to considerable danger, and that 
on the hypothesis descent ought to have been negatived by natural 
selection, i. e. by the development of ligaments or supporting 
areolar tissue. Incidentally noting that the actual facts prove 
that no such danger exists, it may be observed that the testes of 
a typical mammal are very efficiently screened, not only laterally 
by their position between the broad thighs of the hind limbs, but 
also posteriorly by the tail ; and they are well preserved from 
contact with suiTOunding objects by the elevation of the body 
upon its limbs, the case being otherwise in Phocidge, Cetacea, and 
lower animals. 

The ancestral history and arboreal or other habits of the 
Primates constitute sufficient warrant for the conspicuous exter- 
nality of their testes. That active arboreal habits involve 
impulsiveness of the highest degree, is sufficiently manifest on 
contemplating the movements of any of the ordinary monkeys, 
moi-e especially in the case of the Gibbons. 

Thus in a review of the Mammalia, we encounter a considerable 
mass of evidence testifying to the validity of the theory here 
advocated, and more might be added. In the Monotremata, 
Sirenia, Cetacea, most Edentata, Hyracoidea, Proboscidea, and 
Phocidpe, conditions prevail, or have prevailed, negativing the 
descent of the testes ; and these conditions have either consisted 
of the absence of that type of tei-restrial locomotion which has 
been the sole cause of the ti'ansposition of these organs, or of 
secondary factors which have either negatived the operation of 
the primary agency, or effected a reversion of the pre-existing 
effect of the same. 

Since the tiansposition of the testes is mainly due to their mass, 
and definiteness and concentration of form — their means of 
suspension and physiological separateness fi'om the body merely 
constituting conditions to the transposition — it follows that the 
small and diff"use ovaries of the Mammalia will not respond in 
any degree to the forces incident on the body, the magnitude of 
the strains on an attachment obviously being proportional to the 
mass of the organ attached. But apart from the smallness of 
mass possessed by the ovaries, thei'e exists an important reason 
for their retention at or nea,r the primitive position in the body- 
cavity. This reason is the necessity of the proximity of the ovary 
to the oviducal aperture for the conservation of the ova, the ovary 


and tlie oviduct usually pnssossinGj no aniitoniical connection. 
How inipoitant this is may not only be infeired from a jjriori 
considerations, but also from the discovery of the many structural 
devices ada.pte<l to this end. Thus thei'e occuis the " develop- 
ment of special folds of the peiitoneuni, which practically ensure 
the passage of the ova into the oviduct when they are extrudetl 
from the ovaiies. The oviduct, moreovei', has a large and 
finibi'iated mouth, called in human anatomy the ' morsus diaboli.' 
This almost wraps round the ovary, and thus prevents the ova 
from straying in the wrong direction. Moreo\"er, the ovary itself 
is often so ai-ranged that it can easily be withdrawn into a pocket 
of the peritoneum, from which the obvious exit is by the gajiing 
mouth of the oviduct. This disposition of the generative parts is 
still further modified in a few animals, such as the liat and the 
Kinkajou. In these animals the mouth of the oviduct actually 
opens into the inteiior of a closed chamber which contains the 
ovaiy " {Beddard). Thus there exists ample reason for the 
retentive ligaments usually associated with the mammalian ovary 
(see Appendix). 

Tlie Genitalia and Conditions of Locomotion in Ichthyojjsida 
and Saitro'jjsida. 

If, as we have observed, the conditions of manimaJian loco- 
motion alone subject the oi'ganisation to those concussiv e influences 
wdiich have efiected, among other changes, the transposition of 
the testes ; and if, as we have also seen, only the highest mani- 
festations of terresti'ial activity are capable of producing complete 
testicular descent, then we may be certain that neither in the 
relatively inactive tei-restrial Reptilia, the active aerial Aves, nor 
the aquatic Pisces will a like phenomenon occur. In Heptilia, 
though Ophidia and many Lacertilia are capable of bi-ief spasms 
of great activity, the total impulsiveness is very small and of low 
degree. This is not only due to general passivity, but also to the 
fact that " the body of a reptile is, as it wei-e, slung between its 
lin\bs like the body of an eighteenth centuiy cliaiiot ])etween its 
four wheels," with resulting impei-fection of the " i-elations of the 
limbs to the tiunk from the point of view of a terrestiial 
creatuie" [Beddard). Apart from the absence of the causal 
conditions, the depi'ession of the trunk and consequent liability 
of injury to the testes would in itself have negatived descent 
{cf. Phocidje above). 

In the Anura, the only terrestrial amphibia whic^h concern us, 
adoption of terrestrial habits, resulting in a higher degree of 
impulsiveness, has as visual for its concomitants structural con- 
centration and increased retentivity or transposition. Here the 
testes have assumed a " full oval form, compact and undivided : 
they are situated, as shown in the Frog, on the ventral side of 
the anterior half of the kidneys" [Owen). That the testes 
have slightly descended in correspondence with the saltatory 

332 MR. W. WOODLAND ON THE [Apr. 21, 

progression, is sliown on comparison of two such forms as Triton 
and the Frog (see diagram below, p. 334). If, as we admit, anuran 
progression involves on a small scale those impulses to which we 
attribute the descent of the mammalian testes \ wh}^ have not 
these organs migrated more than they have in tliis instance ? In 
the first place, it may be contended that the total impulsiveness of 
a frog, though quite appreciable, is yet of small value as compared 
with tha,t of a mammal of the same dimensions, say the Common 
Mouse. Secondly, the physiological and structural relations 
obtaining between the testes and the kidneys afford sufficient 
grounds for the special retention of the former, and, moreover, 
the firm attachment of the testis to the kidney is shielded from the 
abruptness of incident strains by the slight mobility of the lattei' 
organ. Finally, in addition to the above considerations, it is 
probable that the descent of the testes would here be prohibited 
in consequence of the depression of the hinder portion of the 
trunk and resulting pi'oximity to the external surface. However, 
that such a cause for retention might be operative, it would 
obviously necessitate transposition to an extent that enabled the 
organs to experience adverse influences, and it is questionable as 
to whether they have travelled so far. 

In the typical Aves, as in Mammalia, the expenditure of energy 
is very great, but in the former, as also in Pisces, aquatic 
Amphibia, &c., the bionomic conditions negative impulsiveness of 
locomotion, and their organisation is not subject to the kinetic 
influences which affect terrestrial animals. 

One noteworthy subject is the large increase in volume of the 
testes during the brief courting-season : thus Owen provides a 
figure illustrating the periodic enlargement of these organs in the 
Common Sparrow, showing that their size ranges from that of a 
pin's head to that of a fair-sized marble, half an inch in diameter; 
and this same periodic increase of volume, though not so marked, 
occurs in sundry orders of the Mammalia, e. g. in Rodents and 
Insectivores. It is easily comprehensible that if an animal's 
activity, however impulsive, were largely restricted to the inter- 
breeding periods, the testes, possessing small mass during that 
time, would exhibit less tendency to displacement ; but as yet it 
is not possible to decide whether this factor has any significance. 

Connected with a consideration of the position of the testes in 
birds, is the important case involved by the habits of the 
Ostriches and their allies. The considerable speed attained by 
these birds, comparable indeed with that of the swiftest mammals, 
must involve a certain amount of iropulsiveness ; and such being 
the case, why have not the testes responded by descent ? For the 
situation of the testes in Ostriches is normally avian, being " p»laced 
above and a little external to the kidneys," which latter structui'cs 

are " elongated, flattened, glandular m.asses tying deeply 

seated, and extending from the posterioi- edge of the diaphragm 

1 " The great Bull-Frog may clear six feet at a leap, and repeat them so rapidly 
as to escape a pursuer, unless chased at a great distance from the water " (Owen). 


to the .interior extremity of the pelvic cavity " [Macalister). It 
is obvious that noii-clescent cannot be ascriljed to the serious risk 
of injury that externality would involve (for, unlike as in the 
maniiuals, the position of the proportionately thin legs and the 
absence of a tail ensure no protection from external influences), 
since such an exjjlanation is negatived by the present position of 
the testes. However, if the several chai'acteristics of the mam- 
malian gallop be called to mind, it is easy to aftbrd an explanation 
of this apparently anomalous instance of tlie non-transposition of 
the testes. For, as I shall show, the bipedal progression of the 
Ostrich does not involve impulsive upheavals of the body to any- 
thing like the same extent as in the gallop of a mammal. The 
great upward impulses associated with the upheavals of the mam- 
malian body are in large part due to the elongation of tlie truidv, 
for the depression of one half of the body is largely accelerated by 
the impetus imparted to it on the sudden elevation of the other 
half. That is, the upheaval of one half of the body x-otates the 
whole lengtli of the trunk about its centre of gravity, and so 
causes tlie other half to descend with greater acceleration tlinn 
it would do under the sole influence of gravity. Plence the 
marked intensity of the contacts of the limbs with the earth, 
which have not only to destroy the downward velocity due 
to gravity, and the additional downward velocity due to this 
lever-like action of the trunk as a whole, but have also to 
impart an upward velocity equal in amount to the two it has 
neutralised. In the Ostrich, on the other hand, there is no elon- 
gated trunk possessing two pairs of limbs, and hence there can be 
none of that lever-action just described. Moreover, the individual 
elevations of the body are small compared with those of mammals, 
owing to the alternate action of the limbs (c/. the Kangaroo, in 
which the action of the limbs is not alternate, and in which the 
elevations are consequently great). And lastly, the gliding 
motion adopted by the Ostrich in running is yet anottier factor 
tending to diminish the intensity of the involved concussions. 
Hence for these three reasons the individual impulsive accelera- 
tions imparted to the body of the Ostrich are very small compared 
with those which exist in the case of mammals ; and therefore it 
can easily be understood how it is that, despite the terrestrial 
locomotion and the perpendicular limbs, the testes have not 
descended in this instance. It is also possible that the posterior 
extension of the sternum and ribs in the struthious trunk, 
leading to tight packing of the viscera, so serves as a support for 
the testes (and other organs), but whether this is the case can 
only be determined by I'eferring to the facts. 

In conclusion, it will not be out of place to here give a brief 
summary of the entire argument concerning the cause of the 
descent of the testes, since it will show how strong is the proba- 
bility of the theory advanced. "We have seen that in mammals 
alone there exists a meclianical cause competent to cft'ect trans- 
position. If any doubt remains in the mind of the reader as to 



[Apr. 21, 

the competency of this alleged cause, contemplation of the actual 
locomotion of a horse or dog, or reference to the works of Marey, 
Pettigrew, and others will soon dispel it. Further, I have 
shown that the degree of transposition of the testes corresponds 

Text-fig. 57. 









Diagram illustrating Testis Descent. 

This diagram, based on a series of measurements, illustrates the positions 
and lengths of the testes relatively to the length of the trunk proper (minus tail) in 
Chipea, Scyllium, Triton, Hana, Lacerta (muraJis), Columha, the common mouse, 
and the cat, all measurements being reduced to one scale. The vertical sides of 
the rectangle represent the length of the animal to which the other measurements 
are proportional, and the dots the approximate positions of the centres of gravity of 
the testes. Such a figure well exhibits the concentration and transposition corre- 
sponding to higher grades of impulsiveness. To fully illustrate the relationship a 
similar diagram drawn up from several hundred dissections is necessary, together 
with the average weight of the testis and intensity of the reactions incurred in 
locomotion, appended in the case of each animal. 


to the intensity of the forces concerned, ami have tluis sup})lied 
the inducfcive evidence required to verify the conckision arrived 
at on a priori grounds. It also been pointed out that 
transposition of tlie testes can, from the nature of the function 
of these oi-gans, the mode of emission of their genital products, 
and the absence of adverse external influences, neither cause 
derangements in the individual economy nor impair the fertility 
of the race ; and that in consequence of a similai- change of 
position in the case of other oi'gans inevitably entailing one or 
other of these penalties, the testes alone have descended. More- 
over, the supei-ioi" density, the disposition of the attachment, and 
the appropriateness lioth as regards volume and tlefiniteness and 
concentration of form for change of position, appertaining to the 
testes, lend greater additional facilities to their transposition than 
in the case of othei- organs not so characterised. Hence we can 
not only show why the testes have descended, but also why they 
alone have thus responded to the incident forces — other organs, 
for one or more of the reasons supplied, requiring to be maintained 
in position l)y the special retentive structures developed to tlia,t 
end. Combining with these facts the additional evidence derived 
from a study of mammalian anatomy, showing that imiDulsive 
locomotion is not alone responsible for the descent of the testes, 
but that it has also been the prime cause of the evolution of the 
diaphragm, the metanephi-ic kidney, and other minoi- anatomical 
features, we see still moi-e clearly " how strong is the pi-obability 
of the theory advanced." 

Other Ulustratious of the Relationshljy between Visceral Con- 
formation and Impulsiveness : the Evoliction of the Diaphragin 
aiid the Metanephric Kidney. 

Althoiigh difiering from the reproductive elements of the body 
as regards their economic i-elations to the rest of the organism, 
the non-repi'oductive organs, possessing like mateiial propei'ties, 
may be expected to portiuy simUai- structui-al chai'acteiistics in 
i-elation to locomotor impulsiveness. A consideration of such will 
not only prove of utility as affoi-ding indirect confirmation of 
the foregoing, but will pi-o\dde opportunity for tendei-ing similar 
explanations in the case of other anatomical featui-es. The con- 
stant intei'i'elations of position obtaining between the various 
oi'gans in the vei-tebi-ate body a.i-e in eveiy case easily accounted 
for, either on mechanical or physiological grounds, and it is 
needless to discuss every oi-gan fi-om these points of view ; it will 
suffice if we treat of two or thi-ee structural phenomena, the 
interprets! tion of which is not quite so self-evident or well 

The complete descent of the testes and the pi'esence of a fully- 
developed diaphragm both being conmion chai-acteristics of the 
Mammalia, it is possil)le that the cause to which the foi'mer is 
attributable may also suffice to explain the evolution of the lattei-, 

Proc. Zool. See— 1903, Vol. I. No. XXII. 22 

336 MR. W. WOODLAND ON THE [-^V^'- ^Ij 

and enquiry confirms the supposition. Though, in the above 
account of mammalian locomotion, most stress has been laid upon 
the impulsive elevations of the body and resulting strains on the 
attachments of organs, yet it must not on that account be inferred 
that on descent of either half of the trunk the shock consequent 
on contact of the limbs with the earth (equal in intensity to the 
elevatory impulse) is negligible. (See above in case of Kangaroo 
in which it is taken into account.) Such shocks have the effect 
of causing those viscera which are closely adherent to the mass of 
the body either to exert a considerable pressure on strvictures 
anteiior to them or to be dislodged from their normal position ; 
and if in either case such a result is to be avoided, organs of 
support must be developed. In mammals, the lungs together with 
the heart occupy the anterior portion of the coelomic cavity, and 
behind these are situated the liver, stomach, and intestines, these 
together constituting a considerable mass. These massive organs,, 
unless prevented, would on each contact of the limbs with the 
earth exert great pressure on the fragile compressible lungs 
immediately anterior to them. Hence, in order to obviate 
ensuing derangements, we find in all mammals, and in many other 
terrestrial animals, a stout partition separating the cavity of the 
liver and gut from the cavity of the lungs — a structure which can 
only have been originated by natural selection. Evidence sup- 
poi'ting the conclusion that the diaphi'agm arose as an adaptation 
to the forward pressure of the liver and gastric mass, is not only to 
be found in the fact that the diaphragm is convex anteriorly, but 
also in that the convexity is, cceteris jmribus, proportional to the 
impulsiveness of the animal's activity and to the mass of the 
liver and gut. In herbivores the mass of the gut is greater than 
in carnivores ; on the other hand, "carnivorous (fat-eating) animals 
generally possess a larger liver than hevhixores " {Wiedersheim), 
so that these respective characters tend to defeat comparison \ 
But in the Perissodactyla we find a fairly large liver (considerably 
larger than in Artiodactyla), a long herbivoran gut, and a high 
degree of impulsiveness, the combination of which on our hypo- 
thesis should be correlated with a higlily convex diaphragm — an 
inference which proves correct. Owen remarks that " in the 
perissodactyle Ungulates, in which the movable ribs are numerou.s 
and continued to near the pelvis, the diaplxragm is also extensive 
and much arched towards the thorax " — indicating that the con- 
vexity of the diaphragm is markedly above the normal. And, 
doubtless, other illustrations are adducible. Additional evidence 
as to the primary function of the diaphragm is, moi'eover, afforded 
by the case of the Struthiones (and allies) wliich have adopted 
the mammalian mode of locomotion. In these birds there exists 
a " well-marked diaphragm forming a pai-tition which divides the 
thoracic cavity into two parts, one posterior and small containing 
the lungs, and the other anterior and large containing the heart 

1 The small capacity of the herbivoran thoracic cavity as compared with that of 
the carnivora must be remembered in this connection. 


and liver. It is a fibi-ous membrane, concaA-e forwainls, -with a 
muscular attachment at eithei* side to the ribs and intei-costal 

tissues, which it joins in about the middle of their course 

The pleural cavity is closed above and below by the fil^rous dia- 
phi-agm becoming blended with the first and last i-ibs. The 
anterior thoracic cavity, which contains the pericardium-coated 
heart in its upper ]jart, entirely independent of the pleui'al 
ca\'ity, is di^•ided into two by ;i dense fibi'ous meinbiune 
which springs from two vertebral crura, much as the human 
diaphragm, and extends above the line to join the sternum along 
the border which articulates witli tlie ribs, leaving the heart entirely 
in fi'ont of it ; its concavity is directed downwai-ds and foi-wai'ds, 
and it is separated from the diaphragm propei' ' )y veiy large aii'-cells. 
The liver is completely separated from the al^dominal cavity by a 
fibrous membrane, so that when the included viscei'a are removed, 
it is not at all bi-ought into view. The mesentery is very dense 
and strong " {Garrod ((• Dancin). A comparison of the struthious 
and mammalian diaphragms afibrds furthei- confirmation. In the 
Ostrich, owing to tlie anomalous position of the avian lungs (their 
close application to the costal skeleton ensui'ing the intactness of 
the air-cell connections), the liver has practically assumed their 
role, as i-egards conformation, in relation to the heart and dia- 
phi-agm. Hence the diaphi'agm is concave anteriorly, consequent 
on the necessity for the retention of the liver. In the mammal, 
on the other hand, the lungs are of large volume and freely sus- 
pended in the ventral portion of the thoi-ax, enveloping the heart. 
If the liver wei'e anterior to the diaphi^igm it would, as before 
remai-ked, during locomotion pei-iodically exei't gi'eat pressui'e on 
the lungs ; hence the diaphragm here becomes the foi-ward sup- 
port of the liver and other alimentary viscera. The envelopment 
of the heart by the liver in the Ostrich possibly affords an illus- 
tration of that alternative to the formation of a diaphragm above 
referred to. 

From these and other considerations, we can more precisely 
intei-pi-et the anterior convexity of the diaphi'agm, which is due 
to two causes : the forward pressure in the median line of the 
large abdominal viscera, and the backward lateivil extension of 
the capacious lungs. Evidence as to the cooperation of this last 
factor is affoixled l^y such cases as that of the Manatee, in which, 
on account of its subaqueous habits, unusuall}' developed organs 
of respiration ai'e leijuired, it also lieing essential that their ex- 
tension should be dorsal in older to ensure the ventral situation 
of the centre of gravity of the liody. Here, as Mivart remarks, 
the diaphragm is so oblique that the thorax " extends backwards 
above the whole length of the abdominal viscera" ; and a similai- 
conformation is found in Cetacea and perhaps other groups. In 
development, the diaphragm is " formed from a couple of septa, 
dorsal and venti-al respectively, which arise independently, and 
are for some time quite di.stinct from each other " [Marshall) : 
from which fact it follows that the part the diaphragm plays in 

o38 ME. W. WOODLAND ON THE [^Pl'- 21, 

connection with respiration cannot have constituted its primitive 
raison cVetre. 

Thus, observing the necessity for the development of the dia- 
phragm, we find that its variations in disposition and contour 
found in the Struthiones and Mammalian orders fully confirm 
the conclusion deduced on a priori grounds. 

Another organ which, though not peculiar to the Mammalia, 
is yet a diagnostic feature of terrestrial vertebi-ates, is the meta- 
nephric kidney, and that there is possibly a relationship between 
teii-estrial modes of locomotion and the evolution of this organ 
I will now endeavour to show. 

It is well established that the metanephros is a development 
of the definitive mesonephi'os which has lost its nephrostomata, 
acquired a separate duct, and become more or less concentrated 
in form. It has been pointed out above that concentration 
of structure is an essential concomitant of that increase of 
impulsiveness which results from the increased activity of the 
animal under conditions which involve marked reactions between 
the body and the medium or substratum. That this is so in the 
case of the kidney, it is only necessary to compare the elongated 
mesonephric bodies of Pisces and aquatic Urodeles with the same 
organs of the teirestrial Amphibia and the metanephric bodies 
of Reptilia, and again to compare these latter with their repre- 
sentatives in Mammalia, in which both concentration of structure 
and impulsiveness of locomotion attain their maxima. With 
i-egard to the loss of nephrostomata, the same relation holds. 
In all Pisces, with the exception of certain Elasmobranchii, the 
mesonephi'ic bodies I'etain connection with the coelom by means 
of the nephrostomial tvibules. Whether the absence of these 
fragile structures in Elasmobranchs (their function with regard 
to the ccelom perhaps being assumed by the abdominal pores) is 
to be attributed to the fact that these fishes either are or are not 
descended from the most active members of their class (and it 
must be remembei'ed that they are surface forms) is not certain, 
but it is possible. In the aquatic Ui-odeles the nephi'ostomes are 
present, but in the terrestrial Anura they have completely lost 
their connection with the coelom (which lattei-, as in Elasmo- 
branchs, has had to discover anothei- means of exit for its waste 
prodvicts). The causal relation between disruption of the nephro- 
stomata and the adoption of a tei-restrial life is here clearly shown 
by the ontogeny. In the development of the frog, the meso- 
nephros is at first in communication with the coelom by means of 
the nephrostomata, but at the period of metamorphosis this con- 
nection is severed, and the kidney, becoming more concentrated, 
finally assumes the definite form of the adult structui'e. ISTeedless 
to say, the metanephros of Reptilia, Aves, aiid Mammalia is 
totally devoid of nephrostomes, that of the last not possessing 
them at any stage of development. The division of the meso- 
nephros into two poi-tions, one coming into I'clation with the 
testes, and the other — the definitive mesonephros — retaining its 


uiinaiy function, is evidently a physiological phenomenon, though 
the ultimate structural separation of the two parts must largely 
he attributed to the influenee of greater activity of the organism. 
Also the secondarv development of the definitive mesonephros is 
indirectly due to increased activity, since the worlc of the excretion 
resultina- from the latter is largely increased. In fact, the whole 
ontogenv of the kidney— the successive developments of the pro-, 
meso""-, and metanephros in space and time, the diflerences obtaining 
between the pro- and mesonephric tulmles. tlie disi.ppearance ot 
the pronephros, &c., &c.— can be directly or indirectly attributed 
to the increasing activity of the organism in the course of phylo- 
genv, as a little thought will show. 

" With regard to the position of the kidney— a l)ody of consider- 
able mass, and in Amniota similarly disposed to the testes— no 
relation between such and the impulsiveness of the animal being 
traceable, it is evident that some other factor must be involved 
aiid enquiry shows that the localisation of the kidney ls correlate.! 
with the total activity of the animal. Excretion being facilitated 
h\ proximitv to a vigorous blood-supply, we find that m active 
animals the "kidney tends to be situated anteriorly, i. e. nearer the 
heart ; on the other hand, close connection with tlie vent to the 
exterior is similarly advantageous, and in inactive animals, such 
as Reptilia and Urodeles, we find the ki<lneys situated at the 
posterior extremity of the coelom. 

Tlie kidneys of mammals are retained in position b>- a circuin- 
ambient development of areolar tissue, which usually contains 
much fat ; support is also contributed by the " tonicity " of the 
muscles of the abdominal wall and by the terminal ribs. 

In conclusion, I must acknowledge my indelitedness for details 
respectinc the habits and anatomy of the mammalian orders to 
the following authors and works :' Flower and Lydekker, Owen. 
Wiedersheim. Beddard and Gegenbaur, Lydekker's 'Royal 
Natural Historv.' 'Mammals' by Yogt & Specht, Schmeil's ' Text- 
book of Zoology,' and Allen's & Lloyd's editions of Jardmes 
' Naturalist's Library.' . 

I also wish to thank Prof. Minchin, Mr. J. T. Cunningham, and 
Mr. H. S. Sheltou for kindly reading through the manuscript and 
for making several suggestions and criticisms. 

In respect to the non-descent of the ovary in the Mammalia, it 
is as well to emphasise one factor ensuring its retention. Assum- 
in<v\hat the females of mammals are approximately as active as 
the males, it is evident that if the mammalian o^nlm had retained 
its volk {i. e. if the mammalian ovai'v possessed the size and mass 
of that of the Sauropsida and Monotreines), the retention of the 
ovary within the body-cavity by the development of special liga- 
ments, &c., would have been, if not impossible, yet extremely 
hazardous. And such being the case, it woiUd obviously benefit 
the race if the potency of one of these U\o factors were decreased. 

340 MR. R. I. POCOCK ON THE [Apr, 21, 

Now, under the conditions of mammalian life — conditions in which 
survival of the fittest attains within its limited sphere of operation 
a maximum degree of efficiency — it is obvious that diminution of 
activity wovild be fatal, the speed of mammals being one of the 
most important conditions to survival in the struggle for existence 
(as is shown by the fact that this trait is so highly developed in 
these animals) ; hence, a less degree of activity being prohibited, 
any decrease in the mass of the ovary would be of service to the 
organism in which it occurred. Natural selection may legitimately 
be svTpposed to operate here, since, although it has been clearly 
demonstrated that the higher the life of the organism the less 
range of application does this principle possess, yet it doubtless 
applies in the case of any feature which is of paramount importance, 
and modifications concerned with the genital structures must 
necessarily possess such importance. It is therefore possible, a,nd 
even probable, that the loss of yolk sufi'ered by the mammalian 
ovary and the alternative adoption of a placental mode of nutiition 
both indirectly result from that same cause of impulsive locomotion 
to which we have traced several other features of mammalian 

It may also be worth while to add that many minor features of 
mammalian anatomy, the significance of which is usually over- 
looked, are only explicable on the assumption that they aie related 
to impulsive locomotion. Instances of these minor structures are : 
the accumulation of fat at the base of the heai't, the fatty cushion 
surrounding the neck of the bladder, the fatty development about 
the kidney already noticed, and the various " suspensory ligaments " 
and other " fixative organs " referred to above, associated with the 
stomach, liver, and other viscera of lara,'e mass. 

5. On the Geographical Distribution of Spiders of the 
Order Mygalomorphse. By E. I. PocoCK, F.Z.S. 

fEeceived March 17, 1903.] 

(Text-figures 58-61.) 

Part I. 

Introductory Remarks upon the Paleontology and the 
Means op Dispersal op Spiders. 

(«) Summary of the Palfeontological History of Spiders, and 
its bearing on the Phenomena of Distribution. 

Owing to the enormous chances against the pi'eservation of 
fossil Spiders in sedimentary rocks, the palseontological histoiy 
of this Order is very imperfect. One or two types have been 
discovered in Carboniferous strata of Europe and North Ameiica 
[Arthrolycosa and Protolycosa), and also a fairly large number of 
specimens from amber and from gypsum and lacustrine deposits 
of Oligocene and Miocene age in those countries. But absolutely 


nothing is known of the forms that inhabited tlie world during 
the enormous lapse of time represented by the Mesozoic stiuta, 
and nothing except inferentially of the types that occupied the 
!southei-n countiies of the world dni-ing Tei-tiary times. 

Spiders of the type that lived in the Carboniferous period 
succeeded in holding their own in Europe until the Oligocene, and 
are represented at the present time by the genus Liphistius, which 
is restricted to the Indo- Malayan are;i of the Oriental Region. 

Apait from the genus Liphistius, aW existing Spiders, including 
the Mygalomorpha^, belong to the group Opisthothehe. Thei'e is no 
evidence that this group existed in the Carboniferous period ; but 
since most of the Oligocene and j\[iocene fossils belong to existing 
families, or sometimes indeed to existing genei-a, it is permissible 
to suppose that the OpisthotheljB originated some time during the 
Mesozoic epoch, and may, in fact, be coeval with the mammalia. 
AMiether any of these hypothetic.d INIesozoic foi-ms sui-vive to the 
pi'esent day, it is quite impossible to say. All that palaeontology 
allows us to infei- is that during the Teitiaiy pei-iod thei-e was 
a rich and vaiied spidei'-population spi-ead over the Northei-n 
hemisphere, containing forms that have undergone but little 
metamoi'phosis since that date. The existence of jMygalomoi-ph?e 
at that time is attested by the discoveiy of one foim referred to 
Mygale in the gypsum-beds at Aix, and of anothei', Eoatypus, 
in the Eocene strata at Garnet Bay in the 'Isle of Wight. But 
since it is impossible to classify these forms with an appi-oach 
to cei'tainty in any of the existing families, theii- only value 
from the geographical standpoint is the evidence they supply 
that the Mygalomorphag had come into being in Tertiaiy times, and 
wei'e living in the ISTorthern hemisphere. 

The imperfections in our knowledge above alluded to pei-mit 
oidy a pi'ovisional acceptance of the theoiies put forwai'd in the 
following pages to explain the distributional phenomena of the 
Mygalomoi-pha?. But all the available evidence, little enough 
though it be, points to the conclusion that the jVIygalomorphse 
and the rest of the Opisthothelse appeared iirst in the jSTortheni 
hemisphere, and spread thence over the southein countries of 
the woi'ld. 

{})) jVIeans of Dispei'sal of Spiders, and the importance of the 
Mygalomorphaj fiom the Geographical standpoint. 

It cannot be claimed that Spiders as a whole ai'e a favourable 
group to study from a geographical point of view ; foi-, although 
exclusively terrestrial when adult, and, like other fliiihtless animals, 
dependent upon continuity of land-sui-faces for migration, a gi'eat 
many species are known to have the power, and the instinct to 
pi;t it in foi'ce, of dispelling themselves over wide areas by 
practising when young the habit of flight, using silk-threads as 
aei'ial floats upon which they may be cai-ried long distances before 
the wind. This phenomenon is well known, and has given rise to 

342 MR. R. I. pococK ON THE [Apr. 21, 

the belief in the existence of a ' gossamer ' spider which is supposed 
to be the caxise of the fine thi-eads which fall from the air and 
carpet the fields with silk at certain times of the year. It is now 
known that the ' gossamer ' sjDidei- is a mythical species, and that 
species of the most divei'se habits belonging to widely difi"erent 
fRmilies are responsible foi' the floating threads. The habit is 
practised alike, and, so far as is known, to an equal extent, by 
snare-spinning forms belonging to the Argiopida3 and Theridiidfe, 
by hunting-spidei'S like the Lycosid?e and Attida?, or by sedentary 
species that lurk in flowers, like the Thomisidre. 

That this method of locomotion may considerably influence the 
distribution of spidei's may be infei'red from the fact that cobwebs 
thrown out in this way, and afibrding suppoit to little spiders, 
have been found at the tops of our highest buildings, and have 
become entangled in the rigging of ships 200 miles from land. 

There ai-e reasons for thinking, however, that the habit is for 
the most part restricted to phanerozoic diurnal species, namely, 
those that hunt their prey or spin their webs in the open ; and 
that cryptozoic forms, that live in burrows or under stones or logs 
of wood, and that are for the most part nocturnal, do not indulge 
in it \ 

Clearly, thei'efore, these cryptozoic groups, in which the restric- 
tions to dispersal are piesumably the same as in other terrestrial 
animals which can neither fly nor swim to any distance, have more 
value for the establishment of geographical areas than those species 
with powers of dispersal analogous to flight. 

Owing to the relatively large size and gTeat weight of the newly 
hatched young of the Mygalomorphas, coupled with the reduction 
in the number of spinning-apjjendages and the greater simiDlicity 
of the silk-glands, it seems pi'obable that aerial sailing is not 
practised to any gi'cat extent by the members of this suborder ^. 
Especially true will this be of the Aviculaiiida?, a family which 
contains the largest spiders known of this or any othei' epoch, 
with newly-born young rivalling or excelling in size the adults of 
many species of the Arachnomorpha?. 

Consideiution of these facts, coupled v/ith the impossibility of 
dealing in detail, in one paper, with the distribution of all the 
genera- of the Araiiese, has led to the selection of the Mygalomcrphse 
as the fittest group to illustrate the geographical distiibution of 
Spiders in general. 

1 Simon states that the Spidei'-fauua of the Sandwich Islands is composed wholly 
of species of the former categoiy, with the exception of some few forms which appear 
to owe their presence in that Archipelago to human agencj- (' Fanna Hawaiiensis,' 
Araneaj, 1902). 

" The young of the only known British representative of this group, namely 
Atypus, one of the smallest types of Mygalom()rph;\% have been seen to scatter over 
small areas Ly this method of" travelling \F. Enock, Tr. Ent. Soc. 1885). 

In this connection it is instructive to remark that Att/pvs has a wider distribution 
than any other known genus of the suborder, ranging from Ireland and Algeria to 
Japan and over the Eastern (P the Western) States of North America, that is to say 
across the Northern hemisphere from the eastern to the western shores of the 


Part II. 


MvGALOMORPHiE, and the evidence thus supplied as to their 
Oriijinal Habitat and the Lines of Migration followed in 

Family Diplurid^. 

The Dipluiidif are the most widely distributed of all groups of 
Mygalomorpha>, heing found practically all over the world to the 
south of about the 4Uth parallel of north latitude. The numerous 
groups, however, into which the genera fall present some featui'es 
in their geographical i-ange of considei-able interest. 

1. Subfamily DiPLURix.E.^The genus BrachytheW is met with 
in the Mediterranean Region, Central Asia, and the Southern 
States of Noi'th Amei'ica. Neai'ly allied to it are Hapalothele fi'om 
Madagascai- ; Brachythelisciis from Natal ; Aname, Ixamatus, 
Chenistonia, iind JJekana from Australia and Tasmani;i; Fujius 
fi'om Centi-al and South America ; Trissothele from Chili ; Lycimis 
fi-om the Argentine. South America is also the home of Trechona, 
Diplura, Uriichtis, Harmonicon, and Melodeits. 

2. Subfamily MACROTHELiXiE. — This subfamily is divisil)le into 
four gi'oups. 

(ci) The Macrothelfe : — Maarothele occurs in Spain, China, 
Burma, Singapoi-e, and Java. Neai-ly i-elated ai'e Phj/xiosduema 
fi'om Transcaspia, Stenygrocerciis fi'om New Caledonia and 
Queensland, and Porrhothele from New Zealand. Ischnothele 
is represented liy species from India, Madagascar, S. it W. Africa, 
and Central and South Ameiica ; and Evagrios is known from 
S. Africa and Central America, whence it extends into the 
Southern States of North America (Idaho). 

(/>) The Hexathelfe contain two genera — Hexathele from New 
Zealand, and Scotincecus fi-om Chili. 

(c) The Atraces contain the ^ewei-A Atrax and Hadronycha, from 
Eastern Australia. 

(fZ) The Masteri;e, comprising Accola and J/^asteria, wliicJi are 
probably identical, occur in Yenezuela, the Philippine Islands, and 

The presence of Brachythele in the Mediterranean and Sonoran 
areas, and of nearly allied forms in South Africa, Mailagascar, and 
all over South America, suggests immigration from the north 
into these countries of the Southern hemisphere. On the other 
hand, the entii'e absence of i-elated types from the ai-ea lying 
between and including India and Austro- Malaysia, and the 
reappearance in Atisti-alia of geneia closel}' allied to Brachythele 

' This genus h;is also been recorded ft'oui S. Africit, Madagascar, S. America, and 
Australia. It is probable, however, that the species referred to it belong to one or 
other of the allied 2:enera from these areas. 



[Apr. 21, 












































as well as to the South- American and Afi-o-Mascarene foi-ms, points 
equally forcibly to the peopling of Australia from either one or 
the otlier, or perliaps both, of the southern continents just 

The remaining geneia of Dipluiinaj, namely, IWchona, Diplura, 
Harmoiiicon, Jlelodeus, and Cruchiis, all of which ai'e more 
specialised tjqjes than BracJu/thele and its allies, probably arose 
within their present ni'ea. of (Hstril)ution. 

A greiit contrast to the distribution of the Diplurinog is presented 
by that of the j\Iaci-otlielin;e. 

The occurrence of Macrothele in Spain, Cliina, Burma and Java; 
of Porrhothele, which is scarcely separal:)le from it genericnlly, in 
New Zealand ; of Phyxioschoima iii the Transcaspian ai-ea ; and of 
its near ally, Stenycjrocercas, in Queensland and Kew Caledonia, 
suggests a southward nngr:itiou of these types from the northern 
provinces of the Old Woi-ld into Australia and New Zealand by 
way of China and Indo-iVIalaysia. Similarly Isclinotheh, a more 
specialised type than Macrothele, pei'haps descended from the north 
by way of India into Madagascai-, South Afi-ica, and crossed 
thence into South Ameiica, where with Evayrus, which is also 
I'epi-esented in Soutli Afiica, it is the only i-epi-esentative of this 
group of Dipkn-ida?. The entii-e absence from the Sonoi-an Region 
of foi-ms related to Macrothele, Evagrus, and Ischnothele, is opposed 
to the supposition that the two last-mentioned genera had a 
noi'thern oiigin in Amei'ica.. 

The Mastei-i:>5 appear to be degenerate f oi-ms of the Maci'othela?. 
They ai'e the smallest of all known Mygalomorphfe, and ai-e 
essentially ci-yptozoic or lucifugous, living in cavei'ns or under old 
decaying vegetation in the dark, damp forests. Hence we can 
only pi-etend to a partial knoAvledge of theii- distiibution, and it 
would be rash to draw deductions fi'om the fact of their having 
been discovei-ed hitherto only in Venezuela, the Philippines, and 

The two genei-a of Ati-aces, Atrax and Hadronyche, confined to 
Australia, appear to be Maci"othelina3 specialised for a fossorial 
life, with Avhich is con-elated certain features impai'ting to them a 
superficial similarity to the Ctenizidte. 

The Hexathela?, resembling the Macrothela' except in the 
retention of an additional pair of spinning-mamilla?, undoubtedly 
a primitive feature, ai-e confined, so far as is knoAvn, to New 
Zealand and Chili. There seems no reason to doubt that they 
passed fi-om one of these countiies to the other by a southei'u 

Family Paratropid.e. 

This family, specialised both in sti'ucture and habits, is repre- 
sente<l by three genera, Paratrojns, Anisaspis, and Anisaspoides, 
confined to the Neotropical Region. Its afiinities are doubtful, 
but some primitive genus of Diplurida? allied to Brachythele must 

346 MB. R. I. PococK ON THE [Apr. 21, 

probably be looked to for its ancestry. There seems no reason to 
doubt that it originated in the area it now occupies. 


The genera of Atypidse, two in number, scarcely pass south of 
the Equatoi'. Atypus, occurring in the Mediteri-anean Region, 
and spreading northwards into Central Europe, beyond the 50th 
parallel of north latitude, is the most northern type of the Mygalo- 
morph^e. It is also met with in Japan, Burma and Java, and in 
North America, where it range